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PROCEEDINGS 



(JENEEAL MEETINaS FOR SCIENTIFIC BUSINESS 



OF THE 



ZOOLOGICAL SOCIETY 

OF LONDON. 
1918, pp. 1-196, 

M'lTH 1 Plate and 32 Text-figures. 




PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN EEaENT'S PARK. 

LONDON: 
MESSRS. LONGMANS, GREEN, AND CO. 

PATERNOSTER ROW. 



LI S T 



OP THE 

COUNCIL AND OFFICERS 

OP THE 

ZOOLOGICAL SOCIETY OF LONDON. 

1918. 



patron. 
His Majesty The King. 

COUNCIL. 
His Gbace The Duke of Bedford, K.G., F.R.S., President. 



The Hon. Cecil Baring, M.A. 

Richard H. Burne, Esq., 
M.A. 

Lt.-Ool. S. Monckton Cope- 
man, M.D., F.R.S. 

Charles Drummond, Esq., 
Treastirer. 

Alfred Ezra, Esq., Vice- 
President. 

Capt. Harold S. Ferguson. 

Capt. Hugh S. Gladstone, 
M.A. 

Sidney F. Harmer, Esq., M. A., 
Sc.D., F.R.S., Vice-President. 

Prof. James P. Hill, D.Sc, 
F.R.S. 

William Huntsman, Esq. 

Sir Edmund G. Loder, Bt., 
Vice-President. 



Prof. Ernest W. MacBridb, 
D.Sc, LL.D., F.R.S., Vice- 
President. 

Col. Sir A. Henry McMahon, 
G.C.M.G., K.C.I.E., G.C.y.O., 
C.S.L 

P. Chalmers Mitchell, Esq., 
M.A., D.Sc, LL.D., F.R.S., 
O.B.E., Secretary. 

Adrian D. W. Pollock, Esq. 

The Lord Queenborough. 

The Marquess of Sligo,F.S. a., 
Vice- President. 

AuBYN Trevor-Battye, Esq., 
M.A. 

Anthony H. Wingfield, Esq. 

A. Smith Woodward, Esq., 
LL.D., F.R.S., Vice-Pre- 
sident. 



i'r^hl 



PRINCIPAL OFFICERS. 



P. Chalmers Mitchell, M.A., D.Sc, LL.D., F.R.S., O.B.E., 

Secretary. 
R. I. PococK, F.R.S., F.L.S., Curator of Mammals and 

Resident Superinte^ident of the Gardens. 
D. Seth-Smith, Curator of Bird.s and Ins])ector of Works. 
Lieut. Edward G. Boulenger, Cttrator of Reptiles. 
Prof. H. Maxwell Lefroy, Ci(,rator of Insects. 
John Barrow, Accountant. 
W. H. Cole, Chief Clerk. 



LIST OF CONTEiNTS. 

1918, pp. 1-196. 



EXHIBITIONS AND NOTICES. 

Fage 
The Secretary. Report on the Additions to the Society's 
Menagerie during the months of November and 
December, 1917 187 

Mr. D. Seth Smith, F.Z.S., Curator of Birds. Exhibition 
of,- and remarks upon, a series of lantern-slides made 
from photographs of Reptiles taken in the Gardens... 187 

Dr. P. Chalmers Mitchell, F.R.S., Secretary to the 
Society. Communication of a letter from Mr. T. E. 
Whitehead on the Wild Dingo 188 

Dr. R. Broom, C.M.Z.S. Exhibition of, and remarks upon, 

a New and a Rare Species of the Golden Mole 189 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of January, 1918 190 

Dr. Smith Woodward, F.R.S., V.P.Z.S. Exhibition of a 

copy of an incised drawing of a hunted deer 191 

Professor Poulton, F.R.S., F.Z.S. Communication of a 
letter from Captain G. D. Hale Carpenter on an 
African Civet attacking Human Beings 191 

Professor E. W. MacBride, M.A., D.Sc, F.R.S., F.Z.S. 
An account, illustrated by lantern-slides, of his 
recent investigations into the development of the 
Sea-Urchin 192 

Mr. D. Seth Smith, F.Z.S., Curator of Birds. Exhibition 
and description of skins of the Hoatzin, illustrated 
by lantern-slides 192 



IV 

Page 

Mr. C. Tate Rega.\, M.A., F.R.S., F.Z.S. Exhibition of 
photographs of an Indo-Pacific Ohsetodont Fish. 
(Text-figure 1 .) 192 

The Secretary. Report on the Additions to the Society's 
Menagerie during the months of Febi'uary and 
March, 1918 '. 193 

Miss L. E. Cheesmax, Assistant Curator of Insects. Exhi- 
bition of specimens of an East-African liomopterous 
insect 194 

Dr. A. S iTH Woodward, F.R.S., V.P.Z.S. EKliibition of, 
and remarks upon, the fossil rostral teeth of Eoj^ristis 
and Pristis 194 

Mr. G. A. BouLEXGKR, F.R.S. Exliil)ition of the head of 

a Congo Fish, Hijdrocyon goliath 195 

Mr. D. Seth Smith, F.Z.S. , Curator of Bii'ds. Exhibition 

of, and remarks upon, a Zulu Head-dress 195 

Dr. P. Chalmers Mitchell, F.R.S., Secretary to the 
Society. Remarks on an advertisement announcing 
Fur Sales in the United States 195 

Professor WooD-J ONES, F.Z.S., Honorary Acting Prosector. 
Exhibition of, and remarks upon, specimens from the 
Prosectorium illustrating the effects of Rickets 196 



PAPERS. 



1. Skull of Eana tigrina Daud. By B. L. Bhatia, M.Sc, 

and Baini Prashad, M.Sc. (Assistant Professors of 
Zoology, Government College, Lahore). (Text- 
figures 1-9.) \ 

2. Description of a ne^^• Snake of the Genus Oligodon 

from Upper Burma. By G. A. Boulenger, F.R.S., 
F.Z.S. (Text-figure 1.) g 

3. Reptiles from the River Tajan (Transcaspia). By 

L. A. Lantz. (Plate I.) ,../, , jj 



V 

Page 

4. On the External Characters of the Lemurs and 

of Tarsius. By R. I. PocoCK, F.R.S. (Text- 
figures l-](i.) 19 

5. A Classification of the Pyralid^, subfamily Hypso- 

TROPiN.E. By Sir George Hampson, Bart,, F.Z.S. ... 55 

6. First Report on the Inheritance of Visible and 

Invisible Characters in Silkworms. By Miss Maude 

L. Cleghokn, F.Z.S., F.L.S., F.E.S 133 

7. Notes on Cetacea stranded on the British Coasts during 

1913-1917. By Sidney F. Harmer, Sc.D., F.R.S., 
F.Z.S., Keeper of Zoology in the British Museum 
(Natural History) 1 47 

8. On the Variation of the Pit- Viper, Lachesis atrox. By 

Miss Joan B. Procter, F.Z.S. (Text-figures 1-5.)... 163 

9. Report on Deaths of Animals in the Gardens in 1917. 

By J. A. Murray, M.D., B.Sc, F.Z.S., Director, 
Imperial Cancer Research Fund, Pathologist to the 
Society 183 

Alphabetical List of Contributors vii 

Index xi 



A LPHABICTICAL LIST 



CONTRIBUTORS, 

With liefer ences to the several Articles contributed by each. 
(1918, pp. 1-196.) 



Pnge 

Bhatia, B. L., M.Sc, and Baini Prashad, M.Sc. 

Skull of Rana tigrina Daud. (Text-figures 1-9.) 1 

BouLENGER, G. A., F.R.S., F.Z.S. 

Description of a new Snake of the Genus Oligodon 
from U*pper Burma. (Text-figure 1 .) 9 

Exhibition of the head of a Congo Fish, Hydrocyon 
goliath ■•.. 195 

Broom, Dr. R., C.M.Z.S. 

Exhibition of, and remarks upon, a ISTew and a Rare 
species of the Golden Mole 189 

Cheesman, Miss L. E., Assistant Curator of Insects. 

Exhibition of specimens of an East-African homo- 
pterous insect 194 

Cleghorn, Miss Maude L., F.Z.S., F.L.S., F.E.S. 

Fii'st Report on the Inheritance of Visible and 
Invisible Characters in Silkworms 133 



Hampson, Sir George, Bart,, F.Z.S. 

A Classification of the Pyralid.e, subfamily Hypso- 
TROPIN^ 55 

Harmer, Sidney F., Sc.D., F.R.S., F.Z.S. 

Notes on Cetacea stranded on the British Coasts 
during 1913-1917 147 

Lantz, L. a. 

Reptiles from the River Tajan (Transcaspia.) (Plate I.) 11 

MacBride, Professor E. W., M.A., D.Sc, F.R.S., F.Z.S. 

An account, illustrated by lantern-slides, of his recent 
investigations into the development of the Sea-Urchin... 192 

Mitchell, P. Chalmers, M.A., D.Sc, LL.D., F.R.S., F.Z.S., 

Secretary to the Society. 

Report on the Additions to the Society's Menagerie 
during the months of November and December, 1917 ... 187 

Communication of a letter from Mr. T. E. Whitehead 
on the Wild Dingo 188 

Report on the Additions to the Society's Menagerie 
during the month of January, 1918 190 

Report on the Additions to the Society's Menagerie 
during the months of February and March, 1918 193 

Remarks on an advertisement announcing Fur Sales 
in the United States 195 

Murray, J. A., M.D., B.Sc, F.Z.S., Pathologist to the 
Society. 
Report on Deaths of Animals in the Gardens in 
1917 183 

PococK, R. I., F.R.S., F.L.S., F.Z.S., Curator of Mammals. 
On the External Cliaracters of the Lemurs and of 
Tarsius. (Text-figures 1-16.)...... 19 



IX 

Page 
PoULTON, Professor, F.R.S., F.Z.S. 

Oommunication of a letter from Captain G. D, Hale 
Oarpenter on an African Oivet attacking Human Beings. 191 

Prashad, Baini, M.Sc. See JBhatia, B. L., M.Sc. 

Procter, Miss Joan B., F.Z.S. 

On the Variation of the Pit- Viper, Lachesis atrox. 
(Text-figures 1-5.) 163 

Regan, C. Tate, M.A., F.R.S., F.Z.S. 

Exhibition of p|iotographs of au Indo Pacific Chseto- 
dont Fish. (Text-figure 1.) ; 192 

Seth Smith, D., F.Z.S., Curator of Birds. 

Exhibition of, and remarks upon, a series of lantern- 
slides made from photographs of Reptiles taken in the 
Gardens 187 

Exhibition of, and remarks upon, a Zulu Head-dress... 195 

Wooii-JoNES, Professor, F.Z.S., Honorary Acting Prosector. 

Exhibition of, and remarks upon, specimens from the 
Prosectorium illustrating the effects of Rickets 196 

Woodward, Dr. Smith, F.R.S., V.P.Z.S. 

Exhibition of a copy of an incised drawing of a hunted 
deer 191 

Exhibition of, and remarks upon, the fossil rostral 
teeth of Eopristis and Fristis 194 



Prog. Zool. Soc— 1918. 



INDEX. 

1918.— Pages 1-196. 



[New names in clarendon type. Systematic references in italics. 
(z.s.L.) indicates additions to the Society's Menagerie.] 



Agama sangimiolenta, 14. 
Alamosa hipunctella, 65. 

pvperatella, 65. 

Anerastia ahlepta, 88. 

acrophaa, 88. 

anmmopsis, 88. 

argosticha, 88. - 

haliora, 88. 

ephestiella, 88. 

— ^ erasmia, 88. . 

horhi, 79. 

pleurochorda, 88. 

xipMmela, 88. 

Aptenodytes patagonica (z. s. h.), 
Atascosa quadricolorella, 104. 
Aurora longipalpella, 106. 
AvBs: 

Head-dress made of plumes of 
progne, 195. 
Axis porcinus (z, s. l."), 194. 



BalcBnoptera acutorostrata, 148. 

horealis, 149. 

phy solus, 151. 

Balearica pavonina (z. s. l.), 193. 
Bandera binotella, 89. 

carneella, 90. 

Proc. Zool. Soc. — 1918, 



194. 



Bandera cupidinella, 90. 

perlepidella, 90. 

suhhifeella, 90. 

virginella, 90. 

Baptotropa, gen. n., 116. 

tncolorella, 116. 

Barheria affinitella, 131 . 
Bematiscus lescbse : exhibited, 189. 
Biajra concimiella, 115. 

rhodinella, 115. 

Bison americanus (z. s. l.), 193. 
Boiga irigouatum, 16. 



Gabnia myronella, 131. 

Calamotropa, gen. n., 91. 

pulverivena, sp. n., 91. 

Chere Calera punctilimhella, 59. 

alhicostella, 62. 

Canis familiaris, communication, 188. 

Cardiielis caniceps (z. s. l.), 187. 

Carpliibis spinicollis (z. s. l.), 191. 

Ceiira albifasciata, sp. n., (53. 

discinotella, 63. 

Chere progne: exhibited, 195. 

Chortonoeca, gen. n., 80. 

■ eiM-ysticha, 81. 

leucocraspia, sp. n., 81. 

No. XIV. 14 



INDEX. 



Chortoncecn /niuimrlld, 8\. 

/iiiiiDnin:^, 81. 

Oiirysocliloris s.-ltiteri : oxhibiterl. 

190. 
Ca'iiochroa callfornieUa, 59. 

illlhella, 58. 

inspergella, 59. 

monomncula, 59. 

Coenotropa, gen. n., 89. 

limitella, sp. n , 89. 

Commotria albinervella, sp. u. 

111. 

arrhabdella, sp. n., 108. 

castaneipars, sp. n., 112. 

crasahcapeUa, 110. 

enervella, sp. n., 112. 

erythrograpta, sp. n.. 109. 

invenustoUa, 108. 

laticostella, 108. 

mesiella, sp. n., 108. 

miosticta, sp. n., 110. 

neurias, sp. n., 109. 

oherthiiri, 108. 

opacella, 108. 

phlebicella, sp. n.. 112. 

plioenicias, sp. n., 110. 

-phijcUdhi. 109. 

prophasella, sp. n., 112. 

rhodochroa, sp. n., HI. 

rhodoneura, sp. n., 1 10. 

rosella, sp. n., 111. 

roscopci/nel/a, 108. 

rufidelineata, sp. n., 110. 

rufimedia, sp. n., 109. 

tripartella, sp. n., 109. 

venosella, sp. u., 111. 

Coraco^isis nigra (z. s. l.). 187. 
Coscoroha coscorvha (z. s. l.), 187. 
Critonia hilgerti, 121. 

liolorhoda, 120. 

leucopleura, sp. n., 120. 

ochraeealiti, 121. 

phaeoneura, sp. n., 119. 

■ pro-melcsna, 120. 

purpureotincta, 120. 

. rhodessa, 121. 

roseistrigella, 120. 

sarcoglauca, 121. 



Critonia sarcoida, sp. n., 121. 

suhcoiiciniK'Ua, 120. 

Ci/giius iiidanocepluilus (z. s. l.), 187. 



Belphinus delphis, 159. 
Bembea venulosella, 64. 
Discofroniia normella, 119. 



Echinocai'diuin cordatum : lantern- 
slide, 192. 

Ematheudes crassinotella, 115. 

euchlytella, 115. 

lentistrigalis, 114. 

paleatclla, 114. 

pseudopuiictella, 114. 

punctella, 114. 

stramineUa, 114. 

tunesiella, 114. 

varicella, 115. 

vitellinella, 115. 

Emmaiocera actinoleuca, sp. n. 
127. 

albicostalis, 131. 

ane.rastica, 129. 

aurifusella, 128. 

bifidella. 128. 

earuatella, 130, 

costella, 130. 

cremoricosta, 130. 

depressella, 128. 

diver sella, 130. 

endopyrella, sp. n., 127. 

eremochroa, sp. n., 130. 

ergthrinclla, 130. 

laminella, 129. 

latilimbella, 129. 

leticocincta, 128. 

longiramclla,, 127. 

lucidicostella, 129. 

monochromella, 130. 

ornafella, 127. 

polychroella, sp. u.. 128. 

pulverealis, 127. 

radiatella, 127. 

sanguifusalis, 128. 

strigicostella, 128. 



INDEX. 



Emmalocera subfasckdeUa, 131. 

trioolorai is, 130. 

umhricositella, 129. 

umbrivittella, 131. 

variegatella , 129. 

Epidauria chionocraspis, sp. n., 
92. 

discella, 92. 

perfasciella, sp. n., 92. 

phxniceella, 92. 

strigosa, 92. 

subcostella, sp. n., 93. 

transversarieUa, 92. 

Eremias intermedia, 14. 

{Mesalind) guttulata, 15. 

-= velox velox, 14. 

Eryx miliaris, 17. 
Ethiotropa, gen. n., 116. 

pyromerella, sp. n., 116. 

Ethology. 

Insecta : Exhibition of specimens 

of au East-African homopterous 

insect, 194. 
Eudocimus ruber (z. s. l.), 191. 
Eiimeces schneideri, 15. 
scutatus, 15. 

Felis cbaus (z. s. l.), 194. 
Foiulunkia translucidella, 131. 
Fossifrontia leuconem-ella, 107. 

Gennseus swinlaoii (z. s. l.), 191. 
Geographical: 

Insecta : An East- African homo- 
pterous insect from Nairobi, 194. 
Mammalia: On Cetacea stranded on 

the British Coasts, 147. 
Pisces : Head of Hydrocyon goliath 

from the Congo, 195. 
Reptilia: Reptiles from the Rivei- 
Tajan, 11. 
Globicephala melcEna, 157. 
Gc/rgon taurinus (z. s. l.), 187. 
Grampus griseus, 157. 
Gyranodactylus caspius, 14. 

microlepis, .sp. n. (P). I. fig. 1), 

11. 



Halichoerus grypus (z. s. l.), 187. 
Holacanthns semicirculiitus (Fig. 1) 

photograph, 192. 
Hosidia ochrineurella, 64. 
Hydrocyon goli;ith ; exhibited, 195. 
Hyperoodon roairatm, 153. 
Hypsoij-opa acnidias, 77. 

admnbratella, 73. 

albivenalis, 'iQ. 

approximella, 77. 

biparfella, 78. 

biscrensis, sp. n., 69. 

castella, 78. 

chionorhabda, sp. n., 70. 

contrast ella, 68. 

cremoricosta, sp. u., 74. 

diaphasa, sp. n.. 75. 

dyseimata, 75. 

endorhoda, sp. n, 76. 

■ euryzona, 75. 

fuscostrigeUa, 68. 

fusifasciata, sp. n., 71. 

graptophlebia, sp. n., 76. 

heterocerella, 68. 

icasmopis, lb. 

ichorella, 72. 

illectalis, 69. 

infumatella, 71. 

laropis, 78. 

latcrcidella, 74. 

■ leucocraspis, sp. n., 77. 

leucopldebiella, 11 . 

limbella, 71. 

r luteicosteUa, 70. 

monostidza, sp. n., 76. 

— — neurica, 78. 

niphopleura, 75. 

niveicosta, sp n., 74. 

ochricostella, sp. n., 70. 

ocraceella, sp. n., 69. 

papuasella, 73. 

'paucipunctella, 72. 

periphasa, sp. n., 74. 

pervittella, sp. n., 68. 

phyrdes, 73. 

poly actinia, 76. 

polystictella, sp. n., 77. 

punctinervella, sp. n., 72. 



Hypsotropa purpurella, sp. ii.,71. 

pitt^illclla, 7o. 

'psaiKdtJicUa, 7(). 

quadripunciella, 70. 

ranudoxclla , 76. 

rhodochroella, sp. n., 6'J. 

vliododicha, 77. 

rosescens, sp. n.. 75. 

sabuletdla, 73. 

sceleteUa, 72. 

semirosella, 69. 

lioUpunctella, 72. 

stereosticha, 75. 

subcostella, sj). n., 68. 

si/riaceUa, 72. 

tenuicostella, 69. 

tenuinervella, 73. 

tripar talis, sp. n.. 74. 

will pun dell a, 71. 

verthewisteini, 72. 

" vidneratella, 72. 

zophopleura, 71. 



Inseita. 

On the Pyralida', sublaniiJj Hjp.so- 
tropiniB : systematic, 55 ; On visible 
and invisible characters in Silk- 
worms : structure, 133 ; On an 
East-Af]-ican bomopterous insect, 
194. 

Ityraea nigrocincta : exbibited, 194. 



Lagenurhynchua aculm, 158. 

cilhiroslris, ir.8. 

Laurentia albivenella, sjj. n., 90. 

iiiclarella, 91. 

Leotropa, gen. n.. 64. 

papuanensis, .«p. n., 65. 

phoenicias, sp. n., 64. 

sarcina, sp. n., 64. 

Lophophorus inipeyanus (z. s. l.), 191. 
Lutra lutra (z. s. l.), 191. 



Mahiiia sepleintccniata, 15. 



Mammalia. 

Skull of liana liqriiia: structure, 1; 
External Cluiraclers of the Lemurs 
and of Tarsius: structure, 19; On 
Oetacea sti-anded on tlie Bi'itisb 
Coasts: structure, 147; Couimuni- 
cation of letter on the Wild Dingo, 
188 ; On a new species of the 
Golden Mole ; structure, 189 ; On 
the rare Golden Mole, 190 : Copy 
of drawing of a hunted Deer, 191 ; 
On an African Civet attacking 
Human Beings, 191 : On Fur Sales 
in the United States, 195. 

Martia arisonella, 118. 

Megalophota, gen. n., 117. 

leonella, sp. n., 118. 

Mcuu/hia nanella, 61. 

Mesodiphlehia cimsiveiiia, 62. 
deliquella, 62. 

ochraceella, sp. n., 62. 

ronella, 62. 

stricticostella, 62. 

Meso2ilod,on hidens, 155. 

Metacrateria, gen. n., 79. 

melallactis, 79. 

miasticta, sp. n , 79. 

perirrorella, sp. n., 79. 

ptdverulella, 79. 

Monocienocera hrachiella, 122. 

leticania, 122. 

Morphology. See Stkucture. 



Nasua narica (z, s. l.), 190. 

Nidi-ix tessellata, 17. 

Navasota chionophlebia, sp. n., 

(;6. 

discipunctella, sp. n., 67. 

hgemaphceella, sp. n., 66. 

hebeteUa, 66. 

leuconeurella, sp. n., 66. 

myrioleiia, (17. 

persectella, sp. n., 66. 

syriggia, sp. u., 67. 

Oligodon hamptoni, sp. n. (Fig. 1), 



INDEX. 



XV 



Ollia hovo'ponerella, 104. 

parvella, 104. 

santariteJla, 104. 

Orcinus orca, 15H. 

Osacia lineolella, \'1?>. 

Oxyrhopns cloelia : Inntern-slide, 188. 



P A T II o I. o c; Y. 

Animals in tlie Society's Gardens, 
183. 

Patna brunneicostella, sp. n., 117. 

eborieostclla, 117. 

venatella, sp. n., 117. 

Pectinigeria piDiiponereUa, 104. 

. violodU, 104. 

Peoria albideVa, 78. 

Phocwna 2ihoccena , 159. 

Phcenicopterns chilensis (z. s. l.), 187. 

^ roseus (z. s. h.), 187. 

Physeter catodon, 151. 

Pisces. 

On the development of the Sea- 
Urchin, 192; A Ohsetodont Fish 
with Arabic characters, 192 ; On 
fossil rostral teeth of Eopristis and 
Pristis, 194 ; Exhibition of head 
of a Hydrocyon goliath, 195. 

Polyocha achromatella, sp. n., 
126. 

ciner'ella, 125. , 

detritella, 126. 

— ^ — jlagrantella, 125. 

foioearti, 126. 

fuscicostella, sp. n., 126. 

gensanalis, 125. 

leucopleurella, 124. 

neuropferella, 126. 

plinthochroa, sp. n., 124. 

rhodesim, 131. 

saiiguinariella, 125. 

stipella, 125. 

strigivenella, 125. 

venosa, 125. 

vesculella, 125. 

Polyplectruui chinquis (z. s. l.), 191. 

Poujadia cyttarella, 104. 
leuconeura, 104. 



Poujadia pimella, 104. 
Prinanerastia incarnatella, 80. 

lactealis, 80. 

lotella, 80. 

■ nitid/icosteUa, 80. 

Prophtasia amplichea, sp. n., 105. 

bistriatella, 106. 

epiteuyis, sp. n., 105. 

glaucophssa, sp. n., 105. 

platgcerella, 104. 

pyrostrota, sp. n., 106. 

sphalmatella, sp. n., 106. 



Eagonotia dotalis, 124. 
Raphimetopiis, gen. n., 78. 

abhdella, 78. 

spill ifrontella, 78. 

Reptilia : 

On a new Snake of the Genus 
Oligodon : systematic, 9 ; Reptiles 
frjni the River Tajan : systematic, 
11; On the variation of the Pit- 
Viper : structure, 163 ; Lantern- 
slides made from photographs of 
Reptiles, 188. 

Rhinaphe apofomella, 82. 

approximella, sp. n., 82. 

biseriella, 85. 

brunneovittella, 85. 

castanealis, 83. 

celsella, 83. 

conspersella, 86. 

ella, 86. 

enervella, 84. 

endonephele, sp. n., 87. 

fl.avescentella, 84. 

furmmacula, sp. n., 87. 

haploschema, 84. 

hemirhodella, 85. 

holophaa, 83. 

ignetincta, sp. n., 87. 

infumella, 86. 

lateritiella, sp. n., 86. 

leucotasniella, 85. 

lotricella, 85. 

mictochroella, 87., 

neesimella, 83. 



BhinajjJic iiu/ricoskdis, 83. 

pall klicosl a , 84. 

phaeostrotella, s|). n , 85. 

'pli)tthinri, s;}. 

sangirensis, sp. n., 85. 

secboldi, 87. 

separatella, 84. 

signicoUis. 87. 

stictella, 84. 

syssema, 86. 

- talieUa, 86. 

vectiferella, 83. 

venella, 83. 

venilineella, sp. n.. 86. 

virginella, 84. 

Rhodochrysa superheUa. 89. 



Sahor mania pia, 131. 
Sab'orma forcipella, 122. 

papuacola, sp. n., 123. 

vicina, 123. 

Saluria albivenella, 99. 

anchridis, 98. 

arciicostella, 100. 

ardiferella, 99. 

hreviculella, 101. 

callirhoda, 103. 

cancelliella, 102. 

carnescens, sp. n., 94. 

claricostella, 95. 

ctenucha, 94. 

desertella, sp. n., 97. 

devylderi, 100. 

dichroella, 101. 

distictella, sp. n., 101. 

ensiferella, 102. 

erodeJla, 93. 

flammella, 94. 

flavicosta, sp. n., 96. 

flavipurpurella, sp. n., 98. 

floscella, 96. 

furvella., 100. 

gemmatella, 100. 

Cjlareoaella, 9.'<. 

grammivena, sp. n., 99. 

hemiphceaUs, 94. 

holochroa, 102. 



.Saluria ivficifa. 9(>. 

insignificella, sp. n., 103. 

interpunctella, sp. n., 103. 

lentistrigella, sp. n., 97. 

macrella, 100. 

^ — maculivittdla,, 102. 

magnesiella, 103. 

mesomelanella, sp. n., 103. 

minuteUa, 95. 

muswella, 98. 

neotomella, 101. 

neuricella, sp. n., 98. 

nigritella. 99. 

nilgiriensis, sp. n., 95, 

ocliridorsella, 97. 

opificella, 101. 

ostreella, 102. 

'paranensis, 98. 

paucigraphella, 94. 

pectigerella, 102. 

pleurostichu, 98. 

proleucella, sp. n., 99. 

■ psammatella, sp. n.. 94. 

pulverosa, 102. 

rhodophsea, sp. n., 100. 

rosella, 97. 

rufella, sp. n., 101. 

semirosella, sp, n., 94. 

sepicosfella, 95. 

■ spurcella, dC>. 

stictella, sp. n., 96. 

stictophora, sp. n., 96. 

suhcarjiella, 100, 

subcostella, sp. n., 100. 

tenuicosta, sp. n., 98, 

tetradella, 102. 

tripartella, 103. 

varicosella, 99. 

verecimdella, 97, 

ScJienectadia merilesella, 131, 
Schlegelm wiUoni (z. s. l.), 187. 
Siboga albimediella, sp. n., 113. 

dialeucella, sp. n., 113. 

fnhella, 113. 

zeavora, sp. n., 113. 

Statina Idfasciella, 60. 

cashmiraiis, 60. 

gaudiella, 60. 



INDEX. 



XVU 



Statina punctilineella, 60. 

rhodohafhella, 6^. 

roseotincteUa, 60. 

rosinella, sp. n., 60. 

Structure. 

Insecta : On visible and invisible 

characters in Silkworms, 133. 
Mammalia : Skull of Bana tigrina, 1 ; 
External characters of the Lemurs 
and of Tarsius, 19 ; On Cetacea 
stranded on the British Coasts, 
147 ; On a new species of the 
Golden Mole, 189. 
Ekptilta : On the variation of the 
Pit-Viper, 163. 

^udania siihcostella, 88. 



Tampa dimediatella, 63. 
Taphrometopon lineolatum, 16. 



Tetrapterjx paradisea (z. s. l.), 193. 
Tinerastia discipunctella, 61. 

fissirella, 61. 

Tursiops tnmcatus, 157. 



Variaticn: 

Reptilia : On the variation of the 

Pit-Viper, 163. 
Pisces : A Chsetodont Fish with 
Arabic characters ; On fossil ros- 
tral teeth of Eopristis and Pristis, 
194. 
Viper a lebitina, 15. . 



Zamenis diadema, 16. 

rhodorahchis, 16. 

Ziphius cavirosfris, 154. 



PRINTED liY TAYLOR ANl) FRANCIS, UED LroN COURT, FI.KKT STIil.KT. 



•""- I IT'II 




PARTS 1. OC iJL-»vr-conoi 5^^8©^ 

CONTAINING Pages 1 to 196; with 1 Plate 
AND 32 Text-figures. 



AUGUST 1918. 



PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT'S PARK. 

LONDON : 
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PATERNOSTER ROW. 



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PROCEEDINGS 

OF THE 

aENERiL MEETmaS FOR SCIENTIFIC BUSINESS 

OF THE 

ZOOLOGICAL SOCIETY 

OF LONDON. 
1918. 



&3 [Price Twelve Shillings.] ^^ 



LIST OF CONTENTS. 

1918, Parts I. & II. (pp. 1-196) 



EXHIBITIONS AND NOTICES. 

The Secretary. Eeport on the Additions to the Society's Menagerie during the months 

of November and December, 1917 187 

Mr. D. Seth Smith, F.Z.S., Curator of Birds. Exhibition of, and remarks upon, a series 

of lantern-slides made from photographs of Reptiles taken in the Gardens 187 

Dr. P. Chalmers Mitchell, F.R.S., Secretary to the Society. Communication of a letter 

from Mr. T. E. Whitehead on the Wild Dingo 188 

Dr. R. Broom, C.M.Z.S. Exhibition of, and remarks upon, a New and a Rare Species of 

ithe Golden Mole 189 

The Secretary. Report on the Additions to the Society's Menagerie during the month 

of January, 1918 190 

Dr. Smith Woodward, P.R.S., V.P.Z.S. Exhibition of a copy of an incised drawing of a 

hunted deer , 191 

Professor Poulton, P.R.S., E.Z.S. Communication of a letter from Captain G. D. Hale 

Carpenter on an African Civet attacking Human Beings 191 

Professor E. W. MacBridb, M.A., D.Sc, F.R.S., E.Z.S. An account, illustrated by 

lantern-slides, of his recent investigations into the development of the Sea- Urchin. . 192 

Mr. D. Seth Smith, F.Z.S., Curator of Birds. Exhibition and description of skins of the 

Hoatzin, illustrated by lantern-slides 192 

Mr. C. Tate Regan, M.A., F.R.S., F.Z.S. Exhibition of photographs of an Indo Pacific 

Chffitodont Fish. (Text-figure 1.) 192 

The Secretary, Report on the Additions to the Society's Menagerie during the month 

of February, 1918 I93 

Miss L. E. Chbesman, Assistant Curator of Insects. Exhibition of specimens of an East- 
African homopterous insect I94 

Dr. A. Smith Woodward, F.R.S., V.P.Z.S. Exhibition of, and remarks upon, the fossil 

rostral teeth of Eopristis and Pristis I94 



Contents continued on page 3 of Wrapper. 



PROCEEDINGS 



GENERAL MECTINGS FOR SCIENTIFIC BUSINESS 



ZOOLOGICAL SOCIETY OF LONDON 



PAPERS. 



1. Skull o^ Rana tigrina* Daud. By B. L. Bhatia, M.Sc.^ 
and Baini Peashad, M.Sc. (Assistant Professors of 
Zoology, Government College, Lahore)t. 

[Received August 8, 1917 : Read February 5, 1918. 

(Text-figures 1-9.) 
Index. 



Structure 




In a recent paper Nicholls (2) pointed out that Rana tigrina 
Daud., which is generally nsed as a lal)oratory type throughout 
India, differs markedly in several skeletal and other charactei'S 
from the common European forms {R. tem^yoraria and R. escu- 
Iciita), of which a detailed description is generally given in the 
ordinary English text-books. Nicholls had previously (1) pub- 
lished a Note on the ui-ostyle of R. tigrina and several other 
ji.nurous Amphibia, and in the fiist-meutioned paper he deals 
with the vertebral column, the shoulder-girdle and sternum, 
and the tenth spinal nerve ; the skud of R. tigrina, which pre- 
sents no less marked differences, liowever, has not received 
consideration. 

As far back as 1881, Parker (4), whilst working out the mor- 
phology of the batrachian skull, published short descriptions of 

* [Dr. G. A. Bouleuger, F.R.S., informs me that the Lahore Frog has been 
identified by him as belonging to the typical form. — Kd. P. Z. S.] 
t Communicated by Lieut.-Colonel J. Stephenson, D.Sc, I. M.S., F.Z.S. 

Proc. Zool. Soc— 1918, No. I. 1 



I PROFS. B. L. BIIATXA ANl) BAINI PRASIIAU OH THE 

the skulls of several Indian species including R. tigrina, but the 
account of the latter is incomplete and suffers through having 
been drawn up from the studj' of a single specimen. Both the 
published description and the plate are inaccurate in several 
important respects, and therefore it was considered desirable to 
work out in detail the anatomy of the skull of this common 
Indian frog, and to correct the eri*ors which have crept into the 
otherwise excellent account given by him. Foi- this purpose the 
authors have prepared a large series of fresh skulls, and examined 
them both in the wet and the dry condition. This has been 
.supplemented by an examination of the large number of skulls of 
this type which are used for study by the students working in 
the Government College laboratory. 

The Cranium. 

The skull in this frog is veiy much larger than in the two 
common European species. An average-sized adult skull measured 
40 mm. in the antero-posterior and 37 mm. in the transverse 
direction, while one of the largest measured 45 mm. in the longi- 
tudinal and 48 mm. in tlie transvei-se direction. The cranium, 
which is wide behind, narrows somewhat anteriorly. The superior 
surface is markedly arched, its most prominent point being a 
little in front of the occipital i-egion. All the bones associated 
with the cranium, both investing and replacing, show" a marked 
development; thus the original cartilaginous structure has been 
considerably reduced. As remarked by Parker (4), R. tigrina 
presents one of the most perfect examples of Batrachian cranial 
architecture. 

The Bones of the Cranium. 

The Exoccijntal bones (text-figs. 1-4, ^o.) bounding the foramen 
magnum meet each other in the middle line ventrally, and leave 
only a very small V-shaped area of unossified original cartilage 
between their dorsal ends (all that is left uncovered of the 
original tectum synoticiim). The tectum synoticum never reaches 
the superior border of the foramen magnum, and in the skull of 
older specimens this little area als'o becomes ossified. The two 
bones are seen meeting each other and the slight median pro- 
jection on the posterior border of the fronto-parietals. In 
conformity with the great strength and massive proportions of 
this frog, the occipital condyles are large and are well seen in 
both dorsal and ventral views of the skull (text-figs. 1-4). 
Laterally, where the exoccipital meets the prootic it presents a 
prominent bony ridge {processus mastoideus), there being a thin 
strip of cartilage between the two bones in the young specimen 
only. 

The Prootic bodies (text-figs. 1 ik 7, po.) form a considerable 
portion of the roof and anterior wall of the auditory capsule, and 
extend forwards to form a portion of the inner wall of the orbit. 
Dorsally each presents a quadrilateral ai-ea (vide text-fig. 1) 



SKULL OF RANA TIGRINA. 

Text-figure 1. 
prrt 




q po ^o 

Rana tigrina ; dorsal view of skull. 

a.l., aliiiasal process ; c.a., columella auris ; eo., exoccipital ; f., anterior foutaiiellc ; 
f.o., fenestra ovalis ; fp , fronto-parietal ; m., maxilla ; »., nasal ; _p., palatine ; 
p.c, palatal cavtilagre ; p^n., premaxilla ; p.nl., prenasal process ; po., prootic ; 
^.r.. rostral process; p.rA., rliinal process ; ps., pavasphenoid ; ^^., pterj'goid ; 
q., quadrate cartilage ; qj., quadrato-jugal ; s., squamosal ; se., splienethmoid ; 
sm., septo-maxillary ; s.n., septum nasi ; sp., septum dividing the anterior part 
of the sphenethmoid ; st., stapes ; v., vomer. J, II, III, IV, V, VII, IX, X, 
refer to the foramina for the exit of the cranial nerves. 



Text-fisnre 2. 




JX,X 



H. tigrina ; ventral view of skull. 
For explanation of the letters see text-rig. i. 



1* 



4 PROFS. B. L. BHATIA A\U BAIM PRASHAD OX THE 

l>etween the fronto-parietal and exocoipital on tlie inner sule and 
the inwai"d]y directed horizonti\l flange of the squamosal, which 
partly overlaps it, on the outer. Anteriorly the pi-ootic bone 
forms nearly the whole of the anterior wall of the auditory cap- 
sule : a small are:\ on the outer .^ide of tliis anterior wall remains 
cartilaginous. On the inner side this anterior portion curves 
forwards to form the posterior pAi-t of the inner wall of the 
orbit. In this angle is situated the foi-amen for the exit of 
cranial nerves v. to vii. The foramen is completely surrounded 

TexT-liffm-e 3. 




m. 



no V 



YyVE 



S. tigrina ; lateral view of stall. 

Text-fis'iire 4. 




S. tigrina ; posterior aspect of skall. 
For explanation of the letters sec test-fi?. ]. 



by the prootic bone, and is not merely a notch completed below 
by cartilage. The ventn^l and the posterior walls of the auditory 
capsule are cartilaginous. Postero-latei-ally the prootic extends 
to meet the exoccipitals : at the junction of the two is a curved 
irregular depression, the fossa tympanica. which lodges the 
auditory ossicles and in which the foramen ovale is situated. 
The suspensorium of the lower jaw is attached more externally 
to the side of the auditory capsule. Tiie styloid cartilage is 
attached to the outer side of the cartilaginous portion of the 
capsule. 



SKULL OF RAXA nGKL5>A. D 

The Parasphenoid (text-figs. 2 & 6. ps.) is of the typical form, 
but unusually sti-ong. The transvei-se Kmb lies unrler the 
occipito -auditoiy massej«. Its posterior border is deeply concaTe 
and the ends of the transverse limb are considerably wider than 
the jK>i-tion near the middle line. The po^tei-ior median process 
of the longitudinal arm is short and often elegantly pointed. 
The anterior longer longitudinal arm. besifles forming the floor, 
rises up on each side to foi-m a portion of the Literal wall of the 
cranium. About the middle it Ls broader than at the ends. The 
outer edges articulate with the prootics. the cartilaginous poition 
of the cranium, and the sphenethmoid. .The cartilaginous portion 
of the side-wall of the cranium is relatively mucb smaller than 
the anterior .sphenoidal portion. 

Text-figTire 5. 




yp 



B. fiffrina : separated bones from the sknlL 
pm., prp Tna-riTIa ; m- TnaviTIa ; ■., nasal ; fp- fronto-p^uietal : s^ sqnamo^ftL 

The I'ronto-parietals (text-figs. 1. 3. and o./p-) are adequately 
described by Parker (4) in the following words : — 

•• Above, the fronto-parietals form a strong roof with a 
notch in front, the remains of the fi-ontal suture, but are 
whollv coalesceti beyond this : they end behind in two broad 
winsrs which spi-ead over the hinder region of the ci-aninm 
almost to the end. At first hollow in the middle, in the 
postorbital region they develop a sagittal crest, which opens 
out into two tempoi-al wings. The temporal part dips into 
the orbit and then rises over the ear-masses moulding on to 
their sinuosities. The sides are notched, and the end has a 
concave marsin." 



b PROKS. B. L. IJHATIA AND BAINI PRASHAD ON THE 

On removing the fronto-parietal it is seen that there is only 
one anterior fontanelle (vide text-fig. 7, /.), bounded anteriorly 
by a deep notch in the posterior margin of the sphenethmoid 
and posteriorly by the cartilaginous cranium. The posterior 

Text-figure 6. 





-R. tigrina; separated bones from the skull. 
v., vomei' ; ps., paraspheiioid ; qj., qnadrato-jugal ; q., quadrate ; pt., pterygoid.. 

Text-figure 7. 




P^. b 



eo. 



B. tigrina ; dorsal aspect of skull after removal of investing bones. 
For explanation of the letters see text-fig. 1. 

fontanelles usually found in other forms are absent. Parker, in 
his description, stated "that the fontanelles are presumably like- 
those of the lesser kinds," but this is not the case. 



SKULL OF RANA TIGRINA. 



The S])heneihmoid (text-figs. 1, 2, 3, 7, 8, and 9, se.) is more 
strongly developed than in the European species, and extends 
posteriorly almost to the region of the optic foramen. Anteriorly 
it extends on either side so as to form a part of the anterior 
boundary of the orbit {vide text-fig. 8) ; this feature is much 
better marked in the advilt specimens. In the average adult 
specimen its front portion forms more than half the extent of the 
nasal roof, floor, and the middle wall. In a dorsal view a lozenge 
or diamond-shaped area of this bone {vide text-fig. 1) is left 
between the nasals anteriorly and the anterior median notch of 
the fronto-parietals behind. 

Text-fioure 8. Text-fio-vire 9. 





i?. tigrina. 

Text-fig. 8. — Splienethinoid, dorsal aspect after removal of investing bones. 
se., splienethmoid ; p., palatine ; p.c, palatal cartilage. 

Text-fig. 9. — Splienethmoid, anterior aspect. 

I., apartm-e for olfactorj- nerve ; vp-, septum. 

The cartilaginous skeleton of the nose does not call for any 
special remarks, except that in addition to the rhinal process 
(text-fig. 7,j»j.rA.) there is a median prenasal rostrum {p.r.) in 
continuation of the septum nasi {s.n.). A definite septo-maxillary 
(sm.) is present on either side, extending horizontally from the 
anterior wall into the nasal cavity, though Parker (4) regarded 
them as mere ossifications in the nasal cartilage. 

The Bones of the Face. 

The Nasals (text-figs. 1, 3, & 5, rt,.) are large broad-based 
triangular bones, meeting each other in the middle line and 
diverging posteriorly to enclose the diamond-shaped area of the 
splienethmoid referred to above, and to- meet the anterior ends of 
the fronto-parietals. Anteriorly also they extend as far forwards 
as nearly to reach the nasal processes of the premaxillfe. The 
apex or external angle of the triangular bone is drawn out and 
extends outwards to meet the ascending process of the maxilla. 

The Vomers (text-figs. 2 & 6, v.) meet each other in the middle 
line posterioi-ly, but diverge anteriorly leaving a portion of the 



8 



ON THK SKULL Of RAXA TIGRINA. 



floor of the luisal capsule uncovered. The outei- border presents 
two notches, the posterior one l)ounding the posterior nares. 
The vomerine teeth are situated in an oljlique line along the 
posteiior border. 

The Squamosals (text-figs. 1, 3, 4, & 5, s.) are strongl}' deve- 
loped. From the posterior half of the ci-oss-bar is given ofl' a 
horizontal shelf-like process covering the tegmen tympani and 
extending inwards along its posterior border as far as the 
junction of the prootic and the exoccipital. 

The Pterygoids (pt.), Palatines (/>.), Quadrato-jugals {q.j.), 
Maxillce (m.), bones of the lower jaw and the hyoid apparatus 
are of the usual type. An interesting feature, however, is the 
presence of three well-marked depressions on the ventral side of 
the upper jaw in its anterior portion {vide text-fig. 2). The 
middle one of these depressions is between the two premaxillse ; 
it receives a corresponding median projection from the mento- 
meckelians. The two lateral depi-essions are at the junction of 
the premaxilla Avith the maxilla of each side; each accommodates 
a strongly developed projection of the dentary of each side. 

Literature cited. 

1. NiCHOLLS, George E.^A Note on the Urostyle (o.s coccygeum) 

of the Anurous Amphibia. Pi-oceedings of the Zoological 
Society of London, 1915, p. 239. 

2. NiCHOLLs, George E. — Some Notes upon the Anatomy of 

liana tigrina. Proceedings of the Zoological Society of 
London, 1915, p. 603. 

3. Parker, W. K. — On the Structure and Development of the 

Skull of the Common Frog {Rana temporaria L.). Philo- 
sophical Transactions of the Roval Society of London, 
1871, vol. 161, p. 137. 

4. Parker, W. K. — On the Structure and Development of the 

Skull in the Bntrachia,. — Part III. Philosophical Trans- 
actions of the Royal Society of London, 1881, vol. 172, 
p. 1. 



ON A NEW SNAKE OF THE GENUS OLIGOrOxN. 



9 



2. Description of a new Snake of the Genus Oligodon 
from Upper Burma. By G. A. Boulenger, F.R.S., 
F.Z.S. 

(Publislierl by permission of the Trustees of the British Museum.) 

[Received January 1, 1918 : Head February 5, 1918.] 

(Text-figure 1 .) 

Oligodon hamptoni. 

In 1905 I described a new Oligodon* occupying an isolated 
position in the genus, of which two specimens had been obtained 

Text-fisfure 1. 




Oligodon hamptoni, sp. n. X 1^. 

--at Mogok, Upper Burma, by the late Mr. Herbert Hampton. 

* Oligodon lierherti Boulenger, Journ. Bombay N. H. Soc. xvi. 1995, p. 235, 
pl.fii:!. 1. — The species has been rediscovered in Tonkin andnotic?das O.hei'herti, 
vai'. eberhardti, by Pellegrin, Bull. Soc. Zool. France, xxxv. 1910, p. 30. 



10 ox A NKW SNAKE OF THE GENUS OLIGODON. 

Three years later the British Museum received another specimen 
found at the same place by the same collector and which per- 
tains to the same aberrant group, although unquestionably of a 
distinct species. 1 had put aside the specimen in the hope of 
obtaining others ; but as the hope must now be given up, I 
propose to give a description of this handsome and very remark- 
able Snake. 

Nasal undivided ; portion of rostral seen from above a little 
longer than its distance from the frontal, penetrating rather far 
between the praefrontals ; no internasals ; frontal longer than its 
distance from the end of the jsnout, shorter than the pai'ietals ; 
loreal small, longer than deep ; one prae- and one postocular ; 
temporals 1 + 1 ; five upper labials, second and third entering 
the eye ; three or four lower labials in contact with the anterior 
chin-shields, which are longer than the posterior. Scales in 
15 rows. Vent'-als 160; anal divided; subcaudals 32. A broad 
yellow vertebral band, from the nape to the end of the tail, 
between a pair of reddish brown, black-edged dorsal bands of 
about the same width ; sides bluish grey, with two narrower dark 
bi-own bands, the lower interrupted ; head yellow with dark 
brown markings : a spot capping the tip of the snout, a cres- 
centic band from lip to lip through the ey^es and across the 
snout, an elongate spot on the frontal and on the suture between 
the parietals, connected with a large occipital bifid spot which is 
continued as bands along the body, and an oblique band from the 
parietal to the commissure of the mouth and below. Belly red, 
with black bars occupying a whole shield or interrupted and 
alternating ; lower surface of tail uniform red. 

Total length 540 millim. ; tail 70. 

A single male specimen. 

Although not longer than 0. herherti, this is a much heavier 
Snake, which must be regarded for the present as the largest and 
handsomest of all the Oligodons. 



p. Z. S. 1918, LANTZ. PI. I. 




Bale & Danielsson. Ltd. 



LIZARDS FROM TRANSCASPIA. 



ox REPTILES FHOM THE RIVER TAJAN. 1 1 

o. Reptiles from the River Tajan (Traiiscaspia). 
By L. A. Lantz ^. 

[Received February 5, 1918 : Read February 19, 1918.] 
(Plate I.) 

The small collection of reptiles which 1 a,m about to describe 
was made from April to September 1914 by Mr. N. V.^ 
Meriakri, who presented it to the Zoological Museum of the 
Moscow Univei'sity. Prof. (7. A. Kojevnikof was kind enough 
to entrust me with its study. 

This material, although consisting of only 35 specimens, con- 
tains 16 species, one of which appears to l;e new. It thus affords, 
interesting information on the herpetologicnl fauna of the region, 
of the river TaJan, situated at the meeting of the Persian, Afghan,, 
and Transcaspian frontiers. 

1. Gymxodactylus microlepis, sp. n. (PI. I. fig. 1.) 

3 c? . Length of head and body (from snout to vent) 61, 60,. 
and 40 mm_. 

Head oviform, rather depressed. Snout much longer than the 

diameter of the eye. Forehead slightly concave. Ear-opening 

11 11- .L- 1 Ti J.- length of head „ ^_ , 

small, elhptical. Proportion: , — °- — - =0*27 to 

0-30 ; average 0-28. ^^"g*^^ ^^ ^^«<^^ ^"^^ body 

Body modemte, depressed. A well-marked lateral fold. 

. length of fore limb 

Limbs rather long;'. Proportion : ^j -■ — r--^ -, n — r^ =0'4z. 

° ^ length 01 head and body 

, n ^o A ... T-. X- length of hind limb 

to U"43 ; average 0'43. Proportion: i ,- rn — 1~ 

. = 0-61 to 0-64 r average 0-63. l^^S^^ "* ^'^^^'^ '''''^ body 

Tail cyclotetragonal and slightly depressed at the base, becoming 

1- 1 • 1 . 1 ,1 1 T^ j^- length of tail 

cylindrical towards the end. Proportion : - — — -^ — ^^.j . =-; — ^- 

= 1-33 (1 spec.) length of head and body 

Rostral broader than deep, with median cleft above. Nostril 
pierced between the rostral, the first supralabial, and 3 feebly 
swollen na.sals. 10 or 11 supralabials. Scales of snout, forehead, 
and supraocular region polygonal, slightly convex, small in the 
postnasal depression, enlarged in front of the eye. One enlarged 
superciliary. Paiietal and occipital regions covered with small, 
roundish, feebly convex scales, intermixed with larger, more 
convex, or slightly conical ones. On the temple a few large, 
moderately conical, but not keeled tubercles ; the other temporal 
scales small, granular ; in front of the ear-opening 2 or 3 small 
tubercles. 

Mental large, sub-triangular. 7 to 9 infralabials. 3 pairs 
of chin-shields, the first not forming a suture behind the apex 

* Communicated by G. A. Boulenger, F.R.S., F.Z.S. 



12 JIK. L. A. LANXZ ON 

of the mental. Gular scales extremely small, rouudish, scarcely 
imbricate. 

Neck with very small, juxtaposed, granular scales and longi- 
tudinal rows of large, slightly conical, but not keeled scales, 
changing gradually along the back into 12 or 14 longitudinal 
rows of moderate, elongated, trihedral tubercles ; between the 
median rows a row of small tubercles. The other scales of the 
back lai'ger than those of the neck, flat, feebly imbricate ; across 
the middle of the body 62 to 65 dorsal scales (in a transverse line 
passing between the tubercles). 

Venti'al scales small, cycloid, smooth, 3-5 to 38 in a transverse 
row in the middle of the belly, 136 to 144 in the median line 
from the mental to the vent. 

Suprahumeral scales rather small, imbricate, moi'e or less 
rounded, smooth, or indistinctly keeled. Forearm covered with 
scales like the dorsals, and with a few moderately keeled 
tubercles. Infrahumeval scales granular, -juxtaposed. Infra- 
radial scales like the ventrals. 

Suprafemoral scales imbricated, pointed, smooth, or indistinctly 
keeled on the inner side of the thigh ; the other parts of the 
thigh and leg covered with scales like the dorsals, intermixed 
with large, moderately prominent trihedral tubercles. Infra- 
femoral scales large, roundish, imbricate. Male with a continuous 
series of 34 to 39 femoral and prjeanal pores. Infratibial scales 
similar to the ventrals, but a little larger. 

Tail covered above with transverse rows of very large^ mode- 
rately keeled spinose scales, decreasing in size towards the end of 
the tail ; between these rows of large scales about two rows of 
small, imbricate, more or less distinctly keeled ones. On the 
lower side of the tail, except at its base, a single row of enlarged 
ti'ansverse plates. 

Coloration gi'ey above, with more or less distinct darker trans- 
verse bands, which are disposed as follows : — one on the occipital 
region, one on the. neck, 4 to 7 on the back, about 12 on each 
limb, 12 on the tail. Lower parts white. 

Comparative Notes. 

The presence of a series of numerous femoral and prwanal pores 
•show G. microlepis to belong to the gi'oup of G. caspms Eichw. 
It is especially closely allied to G. fedischenkoi Str. and G. longipes 
ISTik., agreeing with the former in proportions and with the 
latter in most characters of scaling. I am greatly indebted to 
Mr. W. A. Lindholm, vpho was so kind as to examine the types 
and other material of G. longipes, which are preserved in the 
Museum of the Petrograd Academy of Science *. Owing to his 
notes, which complete the description given b}^ Nikolskif, I 
am able to state that these two species are quite distinct. 

* Nos. 8809 (3 (?), 8810 (1 (?, 2 ?), 8811 (1 $), from Neh in Eastern Persia, 
18.V.1896, leg. N. A. Zarudny (types); Nos. 9191 (1 ? ), 9193 (1 (?, 4?), 9194 
(1(?) from the country Zirkuh in Eastern Persia, 21. i v. to 6. v. 1898, leg. N. A. 
Zanidny. 

t Ann. Mus. Zfol. Acad. St. Petersb. 1897, p. 313. 



REPriLKS FROM THE RIVJlR TAJAN. 



la- 



Mr. Lindholm measured specimens of G. longipes and obtained' 
the following data : — 

^ (? ' ? 

Length of fore limb a .j, _^ 

Length of head and body ~ 0-48 to 0-51 ; average O'oO 0-49 to Q-ol; average O'oO 

Length of hind limb „,„„„„ „ _, 

T vf—F-y — 1 A^r-T- = 0"69 to 0-73; average 071 0-65 to 070: average 0-68 

Length ot head and bodj' ' » > n 

The comparison with the corresponding data of G. microlepis 
shows the difierence in the length of limbs*. Besides G. lon- 
gilJes has 12 to 15 supralabials, and the first pair of chin-shields 
almost always forming a suture behind the mental ; on one 
specimen only out of 14 these plates are separated by two small 
scales. As to the size of the tubercles of the head, neck, and; 
back, the scaling of the throat and belly, the number of femoral 
and prfeanal pores, the two species seem to agree. 

There is a very interesting gradation in the characters of 
'scaling between the three species G, mia-olepis, G. fedtschenkoi, 
and G. casjnus. G. microlepis has the smallest and the most 
numerous scales, its tubercles are relatively feebly developed ; 
G . fedtschenkoi forms the link between the foregoing and G. cas- 
pius, which has the lai-gest and the least numei-ous scales, and 
very sti'ongly developed tubercles. The following table con- 
tains the most important tlistinctive characters of these three 
species : — 

G. microlepis. G.fedtschenJcoif. G-. casjOiMs J. 

Tubercles of the temple ... moderately conical. conical. trihedral. 

Before the ear-opening "2 or 3 small tubercle.^. 2 or 3 small tubercles. 1 large tubercle. 

On the neck slightly conical scales, moderately prominent very prominent large 

trihedral tubercles. trihedral tubercles. 

Tubercles of the back moderate, prominent, large, . prominent, not very large, very promi- 

not spinose. spinose. nent, often spinose. 

Gular scales '. : extremely small. very small. small. 

Numlter ot scales in a line^ 
between the apex of the | 

mental or the suture of 'f 136 to 144. 123 to 131. 100 to 114. 

the chin-shields and the | 
anal cleft J 

Number of ventral scales") 

across the middle of the [ 35 to 38. 30 to 33 §. 24 to 29. 

belly ) 

Suprafemorals on the inner I small, smooth, or in- moderate, distinctly large, strongly keeled. . 
side of the thigh 3 distinctly keeled. keeled. 

Number of femoral and-; g 27 to 29 11. 

pra^anal pores ..^ ) ' " 



* Nikolski attributes as a distinctive character to G. longipes the greater diameter 
of the eye, which he supposes to be longer than the distance from eye to nostril ; 
with this statement Mr. Lindholm does not agree. 

t Material : 2 $ from Samarkand. 

X Material: 2 $ from Shemakha (Caucasus); 5 spec. {2 $ , 1 $, 2 juv.) from 
Sangatshaly near l?aku (Caucasus) : 3 spec. (1 (5', 2 $ ) from Askhabad (Transcaspia), 
1 $ from Anaii (Transcaspia), and the ^ described here from the river Tajan. 

§ 24 to 34 ventral scales and 30 to 37 pores, according to Nikolski, Fauna of 
Kussia. Reptiles I., Petrogiad 1915, p. 78. 

II The maximum is 30, according to Nikolski, loc. cit. p. 74. 



14 



MR. I.. A. LANTZ OX 



2. Gymnodac'tylus caspius Eichw. (PI. I. fig. 3.) 

1 ? . Length of head and body 65 mm. 9/10 supralabials, 
7/8 infralabials. 14 longitudinal rows of dorsal tubei-cles. 65 
dorsal scales in a transverse row across the middle of the body. 
101 scales in a line between the suture of the chin-shields and 
the vent. 28 ventral scales across the middle of the belly. 

3. Agama saxguinolenta Pall. 

4 specimens, agreeing in every respect with others from 
Transcaspia (Askhabad, Anau, Bairam-Ali). Length of head and 
body 83 mm. ( d" ), 81 nnn. ( d" ), 78 mm. ( $ ), and 36 mm. (juv.). 
14 to 17 supralabials, 15 to 17 infralabials. 43 to 47 galar scales 
•and 73 to 76 ventral scales in a line from mental to vent. 58 to 

64 dorsal and ventral scales round the middle of the body. The 
young differs from the adults in having no spinose scales. 

4. Eremias velox velox Pall. 

2 specimens, entii-ely agreeing with other material from Tians- 
■caspia (Askhabad, Bairam-Ali). Length of head and body 

65 mm. ( J ) and 34 mm. (juv.). 6 to 9 superciliaries. The lai-ge 
supraocular shields completely or almost completely separated by a 
row of granules from the frontal and the postfrontal. Infranasal 
not reaching the rostral. 6 anterior and 3 posterior supralabials, 
6 to 8 infralabials. 5 or 6 chin-shields in the young, the first 3 or 4 
forming a suture. 22 or 23 gular scales in a line between the 
suture of the chin-shields and the collar. 10 plates in the collar. 
50 to 53 dorsal scales across the middle of tlie bodv. 30 trans- 
verse rows of ventral plates, the longest of which consists of 13 
to 15 plates. 20/21 femoral pores. Supracaudal scales strongly 
keeled. 



5. Eremias intermedia Str. 

3 typical specimens. Length of head and body 55 mm. ( $ ), 
37 and 38-5 mm. (juv.). In the $ a granule between the 
pi-iefrontals. 6 to 8 superciliaries. The large supraocular shields 
entirely separated by a row of granules from the frontal and the 
jDOstfroutal*. 6+1-f 2 to 4 supralabials. 7 or 8 infralabials. 
26 or 27 gular scales in a line between the suture of chin-shields 
and the collar. 11 or 12 plates in the collar. 47 to 50 dorsal 
scales across the middle of the body. 29 or 30 ti^ansverse 
rows of ventral plates, the longest of them consisting of 16 to 
18 plates. 13 or 14 femoral pores on each side. 

* I shall show ill a more detailed publication that the subspecies transeaspica 
Nik., vvliich, according to the author, ma}' be distinguished by this character, is 
"identical with the typical E. intermedia. 



REPTlLEci FROM THE RIVKR TAJAN. 15- 

6. Eremias (Mesalina) gutxulata Licbt. 

1 $ . Length of head and body 49 mm. 5 superciliaries. 
Row of superciliary granules beginning behind the 7th super- 
ciliary only. Occipital as large as the interparietal. 4 anterior 
and 3 posterior supralabials. 7 infralabials. 22 gular scales in a 
line between the suture of the chin-shields and the collar. Collar 
free, consisting of 9 plates. '40 dorsal scales across the middle 
•of the body. 28 transverse rows of ventral plates. 10 femoral 
pores on each side. 

7. EUMECES SCHNEIDERI Daud. 

1 spec. Length of head and body 109 mm. ( J) and 63 mm. 
(hgr.). 6 + 1+3 supralabials in the c^ , 5 + 1 + 2 only in the half- 
grown specimen, the middle one being fused with the following 
by forming a very lai-ge subocular. 8 infralabials. 4/5 or 5/6 
nuchal plates. 25 or 26 dorsal and ventral scales round tlie 
middle of the body. 68 scales in a line between the suture of 
the dliin-shields and the anal plates. 

8. EuMECES scutatus Theob. 

3 spec. Length of head and body 122 mm. (cS), 124 and 
116 mm. (both 2 )• Tli© liead-shields offer many anomalies. In 
one 5 the right supranasal is divided ; the other $ has both 
prsefrontals divided into two unequal parts, and two loreals on 
the left side. In the c? the parietals form a long suture behind 
the interparietal.- 4 to 6 superciliaiies. 4 or 5 + 1 + 3 supra- 
labials, the last being very small, 7 infralabials. 3/4 or 4/4 
nuchal plates. 21 dorsal and ventral scales I'ound the middle of 
the body. 78 to 80 scales in a line between the suture of the 
chin-shields and the anal plates. 

9. Mabuia septemt^niata Reuss. 

2 spec. Length of head and body 82 mm. ( c? ) and 90 mm. 
■( 5 ). Supranasals meeting in a point ( d) or separated ( $ ). 
Prsefrontals separated from each other, the internasal forming a 
short suture with the fi-ontal. 4 anterior and 2 posterior supiu- 
labials. 8 infralabials. 34 or 35 dorsal and ventral scales round 
the middle of the body. 70 or 71 scales in a line between the 
suture of the chin-shields and the vent. 

10. YiPERA LEBETIXA L. 

1 spec* Length of head and body 520 mm. Tail incomplete. 
10 supralabiils, 13/14 infralabials. 25 longitudinal rows of 
dorsal scales. 121 ventral plates. 

* It was impossible to ascertain the sex of the snakes, the viscera having heea 
removed. 



16 Mil. L. A. LANTZ ON 

11. BoiGA TRiGOXATUM Hchiieid, 
2 specimens : — 

Length of head and bodj' 510 480 mm. 

Length of tail 115 108 „ 

Number of supralabials 8/9, tlie 3rd, 4th, and 5th enterinp- 

Number of infralabials 11/12 [the orbit. 

2 + 1 + .3 
Temporal shields 2 + 3 rrirrQ 

Number of rows of dorsal scales 21 

Number of ventral plates 222 221 

Number of pairs of iufracaudals ... 86 84 

12. Taphrometopon lineolatum Brandt. 

1 specimen. Length of head and body 790 mm. Length of tail 
375 mm. 8 supvalabials, tlie 4th and 5th entering the orbit.. 
11 infralabials. 2 + 1/2 + 3 temporal shields. 12 rows of dorsal 
scales. 181 ventral plates. 121 pairs of infracaudals. 

13. Zamenis ruodorhachis Jan. 
2 specimens : — 

Length of head and body 715 630 mm. 

Length oftail 265 230 „ 

Number of supralabials 9, the otli and 6th entering the 

Number of infralabials 10 [orbits 

Temporal shields 2 + 5 to 8 irregular ones. 

Number of rows of dorsal scales 19 

Number of ventral plates 227 226 

Number of pairs of iufracaudals ... 121 117 

These specimens have no red stripe along the back. 

14. Zamenis diadema Schleg. 
4 specimens : — 

Length of head and body 870 830 825 770 mm. 

Lengthoftail 215 190 215 (140) „ 

Number of supralaf)ials 11/12 12'13 10/11 12/13 

Number of infralabials 1113 12 13/14 ■ 12 

Number of rows of dorsal scales ... 27 29 27 27 

Number of ventral plates 225 244 217 234 

Numberof pairs of iufracaudals ... 76 83 83 — • 

Behind the puefrontals a row of 3 accessory shields ; in one- 
specimen the median one is fused with the right inter- 
nasal. 3 [i^i) exceptionally 4 (j:^— ^j frenals. 2 prseociilars.. 
Supralabials more oi- less separated from the loreals and entirely 
separated from the orbit by a row of small shields, the first of 
which may reach the postnasal ; there are 2 or 3 shields between 
the suprala^bials and the loreals, 3 to 5 between the former and 
the orbit, a,nd,ifollowing them, 2 or 3 postoculars. Temple 
covered with small irregular shields. 



REPTILES FROM THE RIVER TAJAN. 17 

15, Natrix tessellata Laur. 

3 specimens : — 

Length of head and body 580 530 470 mm. 

Length of tail 155 155 125 „ 

Number of supralabials 8 

Number of iiifralabials 10/11 

Number of prteoculavs 4 2/3 2 

Number of postoculars 4 4 3 

Temporal shields 1+2 + 2 or 3 

Number of rows of dorsal scales 19 

Number of ventral plates 176 170 176 

Number of pairs of infracaudals 70 73 69 

In the specimen having 2 prteoculars and 3 postoculars the 4th 
and 5th supralabials enter the orbit. 

16. Eryx miliaris Pall. 

1 specimen. Length of head and body 355 mm. Length of 
tail 30 mm. Internasals separated by the point of the rostral. 
4 scales between the postnasals, Kound the eye a circle of 13 
small shields, the lowest of them much enlarged and reaching 
the 6th supralabial, the two anterior ones a little enlarged (prae- 
oculars), the others (supraoculars and postoculars) about equal 
in size. Between the supraoculars 5 scales across the head. 

Between the postnasals and the praeoculars 8 (y+I+i) small 

loreal shields, 13 supralabials, the 3rd one being the highest. 
20 infralabials, the first 3 or 4 enlarged, the others very small, 
with larger shields below them. 

EXPLANATION OF PLATE L 

Fig. 1. Gymnodactylus microle^is, sp. n., ^ , River Tajan. 
Fig. 2. G.fedtschenJcoi Sti'., S, Samarkand. 
Fig. 3. G. caspius Eichw., $, River Tajan. 



Piloc. ZooL. See— 1918, No. II. 



THE EXTERNAL CHARACTERS OF THE LEMURS. 19 

4. On the External Characters of the Lemurs and of 
Tarsius. By R. J. Pocock, F.R.S. 

[Received March 5, 1918; Read March 5, 1918.] 
(Text-figures 1-16.) 

Tabxe o? Contents. 

Page 
Introduction 19 

The Muzzle and the Rhinarium 20 

The Ear 22 

The Facial and Carpal Vibrissge 24 

The Glands of the Pore Limb 25 

The Hands and Feet '27 

The Sublingua 36 

• The Anus and its Glands 40 

The External Genitalia of the Male 42 

The External Genitalia of the Female 47 

General Conclusions and Sj'stematic 51 

Introduction. 

The materials upon which this paper is based are mainly the 
lemviroicl Primates which have died in the Zoological Gardens 
during the past ten years or so. Representatives of practically 
all the commonly imported menagerie species of the group have 
passed through my hands in that time, namely, species belonging 
to the genera Chiromys, Chirogaleiis, Lemur, Gcdago, Perodicticus, 
•and Nycticehiis. For the loan of examples of Hemigcdago and 
Tarsius 1 am indebted to Prof. Wood-Jones, the Society's 
Prosector. I am also indebted to Mr. Oldtield Thomas and to 
Prof. J. P. Hill for the chance to examine other specimens of 
Tarsius. I have not, however, been able in all cases to see repre- 
sentatives of both sexes of the species ; and of many admitted 
genera, notably Alicrocebus. Mixocebus, Lejnlemur, and Loris, no 
specimens have come to hand. This applies also to the three 
genera of Indrisidte {Indris, Propithecus, Lichanotus), which, like 
Tarsius, seem to be intolerant of captivity even in their own 
■countries. The external characters of the Indrisidfe, however, 
have been tolerably fully described and illustrated in Milne- 
Edwards and Graudidier's great work on the Fauna of Mada- 
gascar. From this I have freely borrowed. Unfortunately no 
text accompanies the numerous plates on the various species of 
Lemurida^ published in that worlv. Of other treatises dealing 
with the Lemurs on a comprehensive scale the most important 
is the paper by Mivart and Murie (Tr. Zool. Soc. vii. 1872), in 
which some of the external characters of a few diverse types are 
dealt with from the comparative point of view. The rest of 
the bibliography consulted consists mostly of special memoirs on 
particular species, like Owen's paper on Ghiromys, Burmeister's 

2* 



20 MR. R. I. POC'OC'K OX THE EXTKR.NAI. 

on Tars'ms, Van der Hoeven's on Ferodicticus, Huxley's on 
Arctocebus, Beddard's on HcqDcdemitr, and so forth. 

Many of the facts dealt with in the following pages are 
of course well known. Some chai'acters, however, are here 
described, I believe, for the first time ; and I trust that the 
collation of the facts and their comparative treatment may prove 
useful to future students of this gi'oup. 

In the matter of names I have followed the conservative course 
of using Lemur for the species to which it has been by common 
consent assigned in all recent literature, although by the rules 
of nomenclature, it has no right to a place in the Primates 
at all, but belongs by Storr's very definite selection to the 
Dermoptera, Galeojnthecus volans being its type-species. Even if 
that selection be set aside, it appears to me that the " indications" 
of the 10th edition of the ' Systema' show that the species known 
as Loris tarcUgradus is its type. This is clearly a case for the ' 
" Fiat " Committee on Mammalian generic names ; and it is my 
confidence that the Committee will see the wisdom of allowing 
Leimir catta to stand as the type of Lemui\ that induces me 
to retain this generic name in its commonly accepted sense. 
Similarly I have employed Chiromys for the Aye- Aye [mada- 
gascarlensis), although Daubentonia is the correct title for that 
species. The Fiat Committee, I believe, has these names now 
■ under consideration. 

The Muzzle and the Rhinarium. 

I'he bestial aspect of the face of the Lemuroid as compared 
with the Pithecoid Primates is not due to the general elongation 
of the jaws. In this character the Lemurs are surpassed by 
Papio amongst the Pithecoids. It is due to two correlated 
features, namely, the retention of the primitive moist glandular 
rhinarium and the projection of the upjDer jaw supporting it 
beyond the level of the chin. 

The rhinarium is naked to a A^arying extent on its dorsal side 
and also beneath the nostrils laterally and in front. It is con- 
tinued downwards in front as a strip of grooved naked skin 
cleaving the upper lip to its inferior edge. At this point the lip 
is adherent to the gum covering the premaxillse, so that it is 
incapable of protrusion. 

Although the rhinarium is tolerably similar througliout the 
group, one or two variations may be pointed out. In the typical 
Mascarene Lsmurs, including C'hirogaleus (text-fig. 1, A), the labial 
extension of the rhinarium is comparatively short and tlie later- 
ally extended infranarial portions shallow. In Ferodicticus the 
infranarial portion is deeper ; but in Xycticehtis it is not so. In 
Galago crass icaadat us (tex^t-fig. 1, B) the labial extension of the 
rhinarium is a little longer and thinner than in Chirogaleus 
and Lemur. In Hemigalago demidojffi it is remarkably long and 
gradually widens above wliere it passes into the infranarial portion 



CHARACTERS OF THE LEMURS AND TARSIUS. 



21 



■of the rliinavinm ; but in the length and shape of the rhinarium 
G. seveqalensis is intermediate between G. crassicaiidatus and 
//. demicloffi (text-figs. 1, G; 2, B). 

In Ghiromys the i-hinarinm is not so prominent ; the nostrils 
are longer, more slit-like, and separated in front by a narrower 
septmn. The infivanarial portion is very deep towards the middle 
line, reaching almost to the edge of the upper lip and making the 
labial extension of the rhinarium appear very short. In this 
genus also the frenum which ties the lip to the gum between the 

Text figure 1. 











A. Rliinarium of Chirogalous major. 

B. Rhinarium of G-alac/o crassicaiidatus . 

C. Rhinarium of HemigaJago demidoffi. 

D. Bhinarium of Chiromtjs. 

E. Nose of Tarsius. 



points of insertion of the incisor teeth is somewhat longer than in 
"typical Lemurs, so tha,t the lip is capable of being protruded to a 
slightly greater extent in the middle line. It is probable that 
the variations of the rhinarium and upper lip, like many of the 
structural characters in Ghiromys, are correlated with the rodent 
dentition and peculiar method of feeding of this Lemur ; but our 
knowledge of the function of the rhinarium is too imperfect to 
warrant more than a suggestion on this point (text-fig. 1, D). 



22 Mit. i{. I. rocucK U.N tuk external 

I have not been able to examine in a fresh state the rhinarium 
of any species of Indrisidfe. 

The muzzle of I'arsius is very diflerent from that of all Lemurs.. 
The only trace of the rhinarium, if such it can be called, that 
remains is a narrow rim of naked skin surrounding the nostrils, 
which are widely separated as in the Platyrrhine Pithecoid 
Primates. The nose scarcely projects at all, and the muzzle is 
squarely truncated and deep, and the upper jaw hardly overhangs 
the lower, so that in profile view the muzzle has a decidedly 
feline appearance, contrasting markedly with the generally 
canine appearance of that of the true Lemurs. The long upper 
lip is undivided and continuously hairy from side to side across 
the middle line, and its frenum is set higher above the incisor 
teeth, suggesting that the li]3 is susceptible of partial protrusion 
after the manner of the lip of the pithecoid Primates, but to a 
lesser degree (text-figs. 1, E ; 2, A). 

The primitive muzzle of the lemuroid Primates, with its 
rhinarium and adherent upper lip, is associated with the habit of 
drinking by means of lapping. The specialised muzzle of the 
pithecoid Primates with aborted rhinaiium and pi-otrusible 
upper lip is associated with the habit of diinking by means of 
suction. But Tarsius, although more resembling the pithecoids 
in the structure of the muzzle, drinks, it is said, by the lapping 
method *. This fact is full of interest* in connection Avith the 
view, supported hy other considerations, that Tarsius links the- 
Lemuroids and Pithecoids together. 

The Ear t. 

In the species i-eferred to Lemur the pinna of the ear is 
tolerably uniform in shape and structure. It is small and con- 
cealed to a greater or less extent by its own hairs and those of 
the surrounding aiea of the head. Its superior posterior edge is 
not folded; but the anterior edge of the upper half forms a 
strong ridge overlapping the anterior end of the simple shelf-like- 
svipratragus {plica 'princifalis') and descending below it and 
vanishing in the capsule of the jDinna above and within the 
small lobate tragus. The antitragus is somewhat larger than 
the tragus, sometimes much larger {Lemur catta); and the deep- 
notch between them is approximately on a level with the external 
auditoiy meatus. The flap of the ear behind tlie antitragus is 
marked with a depression, the antei-ior and posterior margins of 
which are respectively the ridge running upwards from the anti- 
tragus and the adjoining postero-infeiior edge of the pinna. 
The supratragus, forming the upper boundary of the capsule of 

* H. Cuming', P. Z. S. 1838, p. 67. Mr. Cuining also makes the interesting 
remark that when any object is put near a Tarsius, the animal " draws up the 
muscles of the face similar to a Monkey and show its ... . teeth." I have never- 
seen Lemurs behave in this vi-ay. 

f Described in several types bj' Mivart and Murie, Tr. Zool. Soc. 1872. 



CHARACTliKS OF THE LEMURS AND TARSIUS. 



23 



the pinna, is set comparatively high up, approximately halfway 
between the tragal notch and the upper edge of the pinna. 

In Chirogaleus (text-fig. 2, C) the ear is similar to that of Lemur, 
but the ear of Microcehus is provided with a much larger pinna 
which, judging from a living example, is ribbed above the capsule 
and capable of being folded as in the Galagos. In the latter, as 
is well known, the pinna is of great size. The portion of it just 
above the capsule is ribbed and grooved and susceptible of 
folding. The supratragus is more expanded and more flap-like 



Text-fie-ure 2. 




A. Head of Tarsius. 

B. Head of Hemiffalai/o demidoffi. 

C. Head of Cldrogaleus major. 
I). Head of Ferodicticva. 

Figures drawn from spirit specimens. All X f . 

than in Lemur and the pouch, probably the homologvie of the- 
bursa of the ear of the Carnivora, is set higher up than in the 
Lemui's. Nycticebus and Perodicticus have the i^inna no larger 
relatively than in Lemur, and it is simplified by the almost total 
suppression of the tragal and antitragal thickenings ; but, as in 
Galago, the pouch is set high up and the supratragus is flap-like 
and valvular (text-fig. 2, D). As Mivart and Murie pointed out, 
thei'e is a small fold of the integument above the supratragus in 
Xycticehus, and a similar but better <leveloped fold is developed 



■24 MR. R. I. POCOCK ON THE EXTERNAL 

in Arctocehus. Huxley cites the presence of this fold as one of 
the features distinguishing this genus from Perodicticus. 

Although the ear of CMromys is relatively as large as in the 
Galagos, it is not ribbed and grooved above the capsxde. The 
supratragus is a thickened i-idge as in Lemur ; but the tragus is 
not an angular projection as in that genus but a simple ridge, 
and the notch between it and the well-developed antitragus is 
comparatively deep and wide. Its lower rim, however, does not 
extend downwards so low as the external auditoiy meatus, the 
portion of the pinna just beyond this meatus being elevated as in 
Carnivora, Ruminants, and many other Mammals. 

The ear of Tarsius is similar in all essential details to that of 
the Galagos, except that the supratragus aiid the antitragus are 
somewhat larger and more valvular (text-fig. 2, A). 

The simplest type of ear in this group, a,nd I suspect the most 
primitive, is that of Lemur and Chirogcdeits, ears with a greatly 
expanded and ribbed pinna and valvular supratragus being 
derivative and specialised structures. According to this view the 
ear of Tarsms is the least primitive of all. It is gradationally 
linked with the ear of Chirogcdeus by the ears of Galago and 
MicrocebiLs. 

The Faded and Carpal Vibrissa}. 

In the development of the facial vibrissfe * the most generalised 
type I have examined is Chirogaleus major, where the mystacial, 
superciliary, genal, and interramal tufts are all well developed 
(text-fig. 2, C). There is a single genal tuft on eadi side set low 
down behind the corner of the mouth. Most of the species 
referred to Lemur resemble Chirogaleus except that the inter- 
ramal tuft is absent ; but in L. variegattts it is usually, if not 
always, retained, although of small size. The full complement of 
tufts is present in Chiromys, but the vibrissas composing them 
are generally shorter than in the typical Lemurs, and in two 
cases the interramal tuft was reduced to a single vibrissa. 

In the Galagos {Galago crassicaudatus, G. senegalensis, and 
Eemigalago demidojfi) the vibrissee are poorly developed as com- 
pared with those of the typical Mascarene Lemurs, more par- 
ticularly Chirogaleus major, with which the Galagos were at one 
time associated. The intei-ramal tuft appears to be invariably 
absent, and the genal tuft is set high iip on the cheek a little 
below and behind the posterior angle of the eye (text-fig. 2, B). 
It resembles in position the upper genal tuft of the typical 
Carnivora, whei-eas in Chirogaleics and Lemur the genal tuft 
resembles in position the lower of the two tufts in that order. 
In Nycticehus and Perodicticios the vibrissas are even less well 
developed than in the Galagos, the genal tuft being suppressed 
in the specimens examined (text-fig. 2, D). The genal and 
intei'ramal tufts are also absent apparently in Tarsius, although 

* P. Z. S. 1914, pp. 889-912. 



CHARACTERS OF THE LEMURS AND TARSIUS. 25 

Burmeister figures some vibrisste in front of the ear. These do 
not, however, correspond in position to tlie genal tuft of the 
Galagos (text-fig. 2, A). 

The prevalence in most orders of Mammalia of the complement 
of tufts descrilied above as occurring in Chirogcdeus suggests that 
the absence of one or more of the tufts is a derivative and not 
a primitive feature. In this I'espect the Lorises, Pottos, and 
Tarsius are more sjjecialised than the Mascarene Lemurs. 

The Carpal vibrissje in Lemurs have been studied by Beddard 
-and Bland-Sutton. Confirming and extending tlieir observations, 
I may add that I have found these tufts of tactile bristles in 
Chirogcdeus major, Ha-palemur grisetts, and in examples of the 
following species of Lenmr, namely, catta, variegatus, macaco, 
mongoz, coronaius, rufiventer, alhifrons, and many of the species, 
subspecies or varieties gi'ouped round the last. They are not 
^always easy to detect in the thick fur, and sometimes appear to 
be wanting ; but in such cases I suspect their absence is due to 
moulting or to artificial removal by rubbing. I have not found 
them in Chiromys, Nycticehiis, Peroclicticus, or Tarsius ; and, since 
Bland-Sutton also noticed their absence in Perodicticus, it may 
be inferred that their absence is characteiistic of the Asiatic and 
African lemuroids*. 

The prevalence of these vibrissas in many orders of Mammals 
suggests that they are a primitive Metatherian and Eutherian 
character, a suijijestion which involves the conclusion that their 
absence in the above-mentioned Lemuroid genera is due to 
svippression and- is a derivative feature. 

The Glands of the Fore Limb. 

In Lemur catta, but in no other species referred to the genus 
Lemur, there is a pecidiar gland on the fore-leg, which was 
figured and described by Biand-Sutton f and also fig-ured by 
Milne-Edwards and Grandidier. A strip of black naked skin 
extends from the palm of the hand over the wrist up the distal 
third of the corresjjonding surface of tiie fore-leg. It ends 
proximally in a smooth elliptical area, which is present even in 
the newly-born young (text-fig. 3, A). In adult males the 
elliptical ai'ea is raised into a. swollen cushion-like pad composed 
of white tissue, fatty in appearance and consistency and covered 
with black skin. On the ulnar side of the pad a large, erect, 
solid horny excrescence is developed (text-fig. 3, B). This varies 
in size apparently with age; its apex is sometimes bifid, but 
generally simple, and it is sometimes present on one limb and 

* I have failed tr> detect the carpal vibi'issse on dried skins of Indris and Propi- 
tJieaus; and in both these genera the interrani'al tnft appears to be absent. Also 
in the one skin of Indris available for examination the genal tvifts are wanting, 
whereas in a skin oi P i-opitheous diadema these tufts are well developed and set 
low down on the cheek as in Lemuridse. 

t Proc. Zool. Soc._1887, pp. 369-372. In 1863 Graj^ pointed out the presence of 
this structure in L. catta and its absence in other species. 



26 



MR. R. I. POCOCK ON THE EXTERNAL 



absent on the other. In females the excrescence is generally 
absent or quite small. Only in rare cases of probably aged 
individuals is it comparable in size to that of the males ; and the- 
elliptical ai-ea is generally flat in the females. 

Text-figure 3. 




A. Fore-arm of Lemur catta, newl^^ born, showing the extension of the naked 

skin from the pahn of the hand and the elliptical glandular area above the- 
wrist. 

B. Section through the glandular area and the horny spur of the fore-arm of an, 

adult male Letnur catta. 

C. Superficial view of the shoulder-gland of an adult male Lenmr catta. 

D. Vertical longitudinal section through the same. 

E. Vertical transverse section through the same. 

F. Section through the shoulder-gland of adult female Lemur catta. 



According to Bland-Sutton the ducts of numerous sweat-glands- 
open upon the surface of the skin of the elliptical area. Secretion, 
no doubt, exudes through the pores ; but I have never succeeded 
in squeezing any visible secretion to the surface b}' pressure. 

The only use I have seen the Lemur make of tlii.s structvire is- 
to apply it to the tail by drawing sinuiltaneously the applied 
wrists along that organ from near the base to the point. 
Possibly by this means the secretion is wiped on the long hairs- 



CHARACTEKS OF THE LEMUKS AND TARSIUS. 2T 

to scent them. But so far as my observations go, this action is^ 
restricted to the male when stimulated to anger at the time 
of rut. 

Also in Lemur catta, but in no other species referred to the 
genus, there is a large gland in the male on the inner side of 
the upper arm near the top of the biceps muscle just below the 
shoulder (text-fig. 3, C). It is a circular or elliptical mound-like 
elevation covered with short hair, except in the centre of the 
summit where there is a small naked area carrying the orifice of 
the gland which is visually shaped like a strongly curved crescent. 
In section the gland is seen to be composed of a thick-walled 
sac, the cavity of which is subdivided by ridges and outgi^owths. 
of the walls, making, in a sense, a multilocular gland. It is the 
partial blocking of the rounded orifice of the gland by one of 
these outgrowths that gives the orifice its crescentic shape. The- 
cavity of the gland is filled with strong-smelling brown sticky 
secretion which under pressure can be squeezed, like a worm, 
from the orifice (text-fig. 3, D, E). 

In the female this gland is not always developed, and when 
present it consists of a small elev^ation covered with normally long 
hair and having no cavity or trace of orifice (text-fig. 3, F). 

The suggestion that these glands are modified mTJmmary glands 
is, I think, erroneous. At all events they coexist with the normal 
pectoral mammary glands. 

It is interesting to recall that Hapcdemar also has a similar 
gland on the shoulder and a somewhat similar gland above the 
wrist. I have, however, seen these only on dried skins, and have 
nothing to add to the descriptions published by Beddard * and 
Bland- Sutton t. 

But there are two points worth attention arising out of the 
facts just mentioned. Despite the development of these glands 
in Loivur catta and Hapalemur griseus^ and in no other species, 
these two Lemurs do not appear to be nearly related. Judging 
from cranial and dental characters, the relationship of Hapalemicr 
griseus is with Prolemicr siimts, and of Lemur caUa with the 
other species usually assigned to the genus Lemur. 

The second point is the coexistence in two otherwise dissimilar 
genera of tico sets of glands, one below the shoulder, the other 
above the wrist. This raises the very important question of 
possible correlation in development between two or more struc- 
tures, a question which opens a very wide field for research in 
zoology. 

The Hands and Feet. 

In the species of Lem,ur the digits of the hand are longish and 
slender, and free from webbing to approximately the same extent 
as in Man. The pollex is the shortest of the series and is sepa- 
rated by a wide space from the second, the base of which it 

* Proc. Zool. Soc. 1884, p. 393 ; and 1891, p. 450. 
t Proc. Zool. Soc. 1887, p. 369. 



■28 



MR. R. I. POCOCK ON THE EXTERNAL 



slightly overlaps when laid forwards. It can be extended at right 
angles to the long axis of the palm, but is not truly ojoposable 
since the large composite palmai- pad — the " ball " of the thumb— 
at its base is almost stationary. The succession of the digits in 
length is 4, 3, 5, 2, but the fourth only exceeds the third slightly, 
and the second and fifth are approximately equal. The palm is 
longish, longer than wide, and passes proximally beyond the base 
of the pollex. Of the four interdigital pads, the first or pollical 
is confluent with the inner proximal (thenar), constituting the 
"ball "of the thumb. The remaining three interdigitals form 
a transverse trilobed cushion-like pad. Behind the fourth or 
external interdigital, the external portion of the palm is occupied 

Text-fie-ure 4. 







A. Foot and B. 'Hand of Lemnr macaco ; X f . 
1, 2, 3, 4, the intermediate pads ; I, II, the proximal pads. 



by the elongated external proximal (hypothennr) pad, which is 
subdivided, the posterior expanded portion lying alongside the 
posterior portion of the "ball "of the pollex, from which it is 
separated by a, groove (text-fig. 4, B). 

In its main features the hand only varies in minor particulars 
in the difierent species examined, namely, L_. catta, macaco, alhi- 
frons, mongoz, variegatus, and coronatus, except that in L. varie- 
gatus the palm is a little wider as compared with its length. 

In the foot the hallux is of great lengtli and thickness, is 
capable of being extended at right angles to the sole, and is 
opposable, the " ball " of the hallux, consisting mainly of the 
large, projecting first or hallucal interdigital pad, being movable 



CHARACTEllS OF TJJE LKMURS AND TARSIUS. 



29- 



and capable of being pressed ngainst the second interdigital pad 
and the ehjngated external proximal pad, closing up the depression 
in the middle of the lower surface of the foot. For the rest, the 
digits and the pads of the foot resemble in a general w{\j those 
of the hand (text-fig. 4, A). 

A point to notice is that in Lemur catta the naked sole is 
extended proximally to the tip of the calcaneum or heel. In the 
other species the heel is covered with hair, the hairy area being 
about one-third the length of the naked area, or a little more. 



Text-figure 5. 





A. Hand and B. Yoot oi Chirogalevs major \ nat, size.j 
C. Hand and D. ¥ooi oi Galacjo senegalensis; nat. size. 



In Chirogaleus major the hands and feet are substantially 
similar to those of Lemur, but the naked palmar and plantar 
surfaces are broader as compared with their length, and the 
individual pads are more sharply difierentiated, and, judgino- by 
their papillary ridges, endowed with gi'eater tactile sensibility -^ 
In the foot the hairy area of the heel is relatively longer, being 



30 



MR. R. I. POCOCK ON THE EXTERNAL 



about two-thirds the length of the naked portion. In the specie 
men examined the third and fourth digits were appr-oximately 
equal both on the hands and feet (text-fig. 5, A, B). 

The hands and feet of the Indrisida?, as figured and described 
by Milne-Edwards and Grandidier, show some inteiesting pecu- 
liarities suggesting more complete adaptation to arboreal life than 
in the Lemurida?. In Propithecus the digits of the hand are 
hardly more webbed than in Lemur, but the palm is narrower, 
■especially posteriorly, where the pollex arises. The pads appear 
to be very little diflerentiated, and the absence of the "ball" of 
the thumb indicates a feeble grip for that digit. In Lichanotus 
(Avahis) the ball of the thumb is better developed, but the palm 
•of the hand is apparently longer than in Propithecus, the second 

Text-fiffure 6. 




A. Foot and B. Hand of CMromys; X f. 

•digit, which is very short, being widely separated from the 
pollex. The third, fourth, and fifth are long and united by 
narrow webbing approximately to the ends of the first phalanges. 
In Indris also the hand is long and slender, with a wide space 
between the long weak pollex and the second digit. The latter, 
however, is not so short as in Lichanotus, and is united to the 
third, as the third is to the fovirth and the foui'th to the fifth, by 
integument permitting but slight divarication of these digits and 
increasing in appearance the elongation of the palm. 

In the feet there is less variation. The hallux is normally 
-elongated, but is slender from base to apex without the muscular 
development seen in other Lemuroidea, and the digits are webbed 



CHARACTERS OF THE LEMURS AND TARSIUS. ' 31 

approximately to the distal ends of the first phalanges. In 
Indris the interdigital webbing is somewhat deeper than in the 
others and extends to an equal distance along all the digits, but 
in Fropithecus and Lichcmotus it is deeper between the third 
and fourth than between the second and third and the fourth 
and fifth. 

Some of the peculiarities of the hand of Chiromys, such as the 
presence of claws and the modification of the third digit, ai-e well 
known (text-fig. 6, B). The palm is narrower than in the true 
Xiemurs. The pollex is not truly opposable, but it is so long that 
when turned forwards it overlaps the palm, as in Man. All the 
other digits are long, even the second and fifth, which are sub- 
equal, exceeding the length of the palm. The third is very long 
and slender ; the metacarpal that supports it projects beyond the 
palm, a unique modification which confers exceptional mobility 
op the digit. Nevertheless this digit is shorter than the fourth, 
which is nearly twice as long as the palm. Although a shallow 
Aveb joins tliese two digits at the base, the hand may be described 
as unwebbed. In correlation with the grasping capacity of the 
large claws, the digital pads are comparatively slightly exjjanded. 

The feet are less modified than the hands and conform to those 
of the Lemur-type, except that the digits are thinner, the hallux 
is a little shorter and weaker, the second, thiixl, fourth, and fifth 
are a little longer and armed with claws correlated with narrower 
pads. The heel is covered with hair as in all Lemurs except 
L. catta (text-fig. 6, A). 

In this genus the unique elongation of the metacarpal of the 
third, digit of the hand and the slenderness of the digit are 
adaptations to feeding. 

In Galago of the crassicaudatus and seiiegalensis types the hand 
is wider than in Chirogcdeiis and its pads still better differentiated 
and more prominent, especially the first, or pollical, intermediate. 
All the nomaal six pads are distinguishable. The internal 
proximal is a small pad situated near the base of the outer border 
of the thumb, and the external proximal is a rounded pad in 
contact with the internal intermediate and not extending back- 
wards to the wrist as in Chirdgcdeics and the true Lemui's. A 
wide space separates it from the internal proximal, which is also 
set nearer the wrist. Similar differences obtain in the feet, 
where all the six pads are clearly defined. In G. crassicaudaPas, 
monteiri and their allies the posterior part of the naked portion 
of the foot, as Beddard pointed out, is covered Avith coarse close- 
set papilhie, and the hairy area of the heel is longer than in 
Chirogaletis majo?', being about as long as tlie naked portion. 
In G. senegaleitsis it is considerably longer (text-figs. 5, C, D ; 
7, A, B). 

In Hemigalago the pads show some interesting differences 
from those of Galago. In the hand the six pads ai-e even more 
sharply defined, and are arranged so as almost to encircle a large 
naked submedian area of wrinkled skin. The two proxiujal pads 



32 



MR. R. I. POCOCK ON THE liXTKRNAL 



are almost in contact in the middle line, the external being elon- 
gated and extending practicnlly from the wi'ist up to the internal 
intermediate. Behind the two proximal pads there is a short 

Text-fisfure 7. 




A. Hand and B. P"oot of Galago crassicaudatiis. 
1-4, the intermediate pads; I, II, the proximal pads. 



area of naked wrinkled skin. The pollex is longer and more 
prehensile than in Gcdayo. The arrangement of pads on the 
foot is similar, but the two proximal pads are relatively much 
smaller than in the hand, and the internal is set farther from the 



CHARACTERS OF THE LEMURS A\D TARSIUS. 



33 



hallucal intermediate and is widely separated from the external. 
In this genus there is no definite heel-like area behind the pads 
as in Galago, the skin ronnd the proximal pads being merely 
soft and creased. The haiiy area of the foot back to the tip 
of the calcaneum is nearly twice as long as the naked area (text- 
fig. 9, A, B). 

Text-fis-ure 8. 







A. Hand and B. Foot of Ni/cticehns ; X |. 
C. Hand and D. Foot of Perodicticus ; X f • 
1-4, the intermediate pads ; I, II, the proximal pads. 



The arrangement and distribution of the pads give a primitive 
stamp to the hands and feet of Hemigalago in the sense that they 
recall very forcibly the pad-development seen in the extremities 
of many Rodents, Insectivores, and Marsupials. 

The hand of Nycticebus " and Perodicticios differs in some 
Proc. Zool. Soc— 1918, No. III. 3 



34 MR. R. I. POCOCK ON THE EXTERNAL 

remarkable particulars from that of the other genera hitherto 
noticed. It is capable of being turned at right angles to the axis 
of the forearm. Tlie pahn is short and broad, and the powerful 
pollex can be set so far backwards that its long axis is practically 
in the same line as that of the palm aiid of the fourth and 
longest digit, thus giving the widest possible span to the extre- 
niit}^ and when the pollex is in this position the internal proximal 
pad, constituting the proximal portion of the "ball" of the 
thumb, lies nearly in a line behind the external proximal pad. 
Tlie pollex, moreover, is incomparably more strongly opposable 
than in any other genus of Lemuroid Primates, and surpasses 
even the thumb of Man in that respect. When brought into 
opposition, the composite ball of the thumb is pressed against the 
second intermediate pad and the external proximal pad. Except 
that the hallux is a little longer and stronger than the pollex, 
the feet conform to the hands in type. In all other genera of 
Primates the two extremities are dissimilar. 

As is well known, the second digit of the hand in Nycticebus is 
dwarfed and in Perodicticus is represented by an excrescence 
upon the second intermediate pad. For the rest the hand of 
Perodicticus is narrower with reference to its span, and the third 
and fifth digits are tied basally by a shallow web to the fourth, 
so that the three cannot be so widely separated as in A^i/ciicebus. 
In the feet of both genera the heel is hair}^ and shorter than the 
naked part of the sole, the second digit is dwarfed, and the third 
and fifth are basally webbed to the foui'th in Perodicticus, free 
in Nycticehus, The nails on both extremities are larger in the 
former than in the latter genus (text-fig. 8, A, B, C, D). 

The published descriptions of the extremities of Arctocehus 
suggest that they differ from those of Perodicticus in having the 
third, fourth, and fifth digits more completely webbed (see P. Z. S. 
1864, pp. 316-317, 319-320). Huxley, however, states that the 
calcaneal tuberosity of the foot is naked and separated from the 
padded portion by a narrow band of hair. But since the heel is 
without exception covered with hair in all the remaining species 
of arboreal * lemuroid Primates, I suspect that the nakedness 
mentioned by Huxley was due to artificial rubbing in the specimen 
he examined. It is not uncommon for the hair to be worn 
o(i' the heels in captive examples of common lemurs (Z. albifrons, 
etc.). 

In the distinctness and separation of the pads and their 
encircling ai-r.angement round a cential palmar ai-ea,, the hand of 
Tarsius recalls that of Hemiyalago ; but there are certain differ- 
ences. The hand and fingers of Tarsius are relatively longer, 
the pollex is smaller, closer to the second digit and not opposable, 
the third digit is the longest of the series, slightly surpassing the 
second and fourtli, which are subequal. The first, or pollical, 

* Lemiir criffa, winch lias the heels iiaked alone: the middle line, lives in 
rocky hills. 



CHARACTERS OF THE LEMURS AND TARSIUS. 



35 



intermediate pad is small and rounded, smallei- than and in con- 
tact with the internal proximal, which is elongated and surpasses 
the external proximal in length. The second intermediate pad 
is markedly larger than the others of tha,t series. Finally, the 
nails are sharply pointed, compressed and convex from base to 
tip, simulating claws, nearly the distal half being free from the 
pad thougli normally resting upon it (text-fig. 9, C). 

Text-figure 9. 




A. Hand and B. Voot oi Heniigalago demidqffi ; nat. size. 
C. Hand and D. Koot of Tarsius ; nat. size. 

1-4, intermediate pads ; I-II, proximal pads. 



In the feet the hairy area back to the heel is shorter than in 
Hemigalago, and the naked padded area and digits longer. The 
plantar pads are peculiar. The hallucal, or fii-st, intermediate is 
large and prominent, and is fused to the internal proximal to 

3* 



36 MR. R. I. POCOCK ON THE EXTERNAL 

form a single long pad which sometimes fuses proximally in the 
middle line with the external proximal, which is itself united to 
the external intermediate, forming a long pad extending along 
the outer edge of the sole of the foot. The second and thii-d 
intermediates are united to form a single large elongated pad, 
broad distally, narrowed proximally, whei'e it terminates in the 
middle of the membranous area of tlie sole. Of digits two to five 
the fourth is the longest as in Lemuroid Primates; but the third 
has an erect claw like the second — a characteristic not found 
in any Lemuroids, but probably primitive and forcibly recalling 
the con-esponding " syndactyle " digits of many Marsupials (text- 
fig. 9, D). 

The Suhlingua. 

There are one or two points to be recorded in connection with 
the suhlingua, an organ which is especially well developed in the 
Lemuroid Primates. Typically it is a thin flat fibrous plate, 
lyrate or leaf -like in form with free apex and free lateral margins. 
It covers a considerable area of the lower surface of the tongue, 
the frenum of which arises from an angular notch in the middle 
of its posterior border. The pointed or truncated apex is ser- 
rated or denticulated to a varying extent, and the underside is 
strengthened by a fine median longitudinal ridge, with frequently 
a smaller ridge close to it on each side, making three in all. 

Beneath the sublingua the floor of the mouth is provided in 
front with a pair of small soft flaps, the frenal lamellce, arising 
at the bottom of the lingual frenum and continued backwards to 
a varying extent as a free nari-ow edge towards the base of the 
tongue. A similar and homologous structure is pr( sent in some 
other Mammals — e. g., Canis, Pteropus, where, as in the Primates, 
it overlies the orifices of the sublingual and submaxillary salivary 
glands. 

In the species usually referred to Lemur the sublingua ex- 
hibits certain structual differences, which examination of a larger 
number of individuals than I have seen may show to have 
systematic value. For instance, in L. variegatus there are 
three comparatively strong denticles at the tip and three corre- 
spondingly strong keels below. In other species the tip is x-ather 
serrulate and the keels weaker. On the_ other hand, I have noticed 
differences in the shape of the sublingua within specific limits. 
In a male of L. coronatit.s^ for example, it was lyrate witli bulging 
postero-lateral margins ; in a female it was evenly attenuated, 
the two margins gradually converging from near the base to the 
tip. In the male again the frenal lamellae formed together a 
broad semicircular flap ; in the female they were narrower, 
forming a nearly parallel-sided flap. In a, male specimen of 
L. cdhifrons alhifrons and in a female of L. a. nigrifrons the 
frenal lamellaj were of the same shape as in the male L. coro- 
natus ; but in examples of L. variegatus, macaco, catta, and mongoz 
the laraellte were narrow and resembled more or less closely those 



CHARACTERS OF THE LEMURS AND TARSIUS. 



37 



of the female L. coronatus mentioned above (text-fig. 10, A-D, G). 
In Chirogaleus major the narrowed sublingiia is apically serrulate 
and the frenal lamellae are narrow as in most Lemurs, The 
sublingua of Hapalemitr was briefly described by Beddard. 

Text-fieure 10. 




A. Lower side of tongue of Lemur coronatus. 

B. Sublingua of another specimen of the aame species raised from the frenal 

lamellae. 

C. Lower side of distal end of tongue of Lemur variegatus with the tip of the 

sublingua showing beyond the frenal lamellae. 

D. Lower side of tongue of the same without the frenal lamellas. 

E. Lower side of tongue of G-alago crassicaudatus. 

P. The same of Nycticehtis with supplementary frenal lamellas. 
G. Side view of tongue of Lemur coronatus. 
H. The same of Galago crassicaudatus, 

si., sublingua ; fl,, frenal lamellae. 



38 MR. R. I. POCOCK ON THE EXTERNAL 

The subliiigua is also present, as Milne-Edwards and Grandidier 
have shown, in the Indrisidse.' It has a single median inferior 
crest and three apical denticles. In Propithecus it hardly differs 
in form from that of the Lemuridse, being broad at the base 
posteriorly, nearly parallel-sided, and attenuated at the apex. 
In Indris its lateral edges ai'e rather strongly convex, the base 
being narrowed like the apex. In Lichanotus it is very short, 
its free lateral edge being apparently only about half as long 
relatively as in Propithecus and Indris. 

In Galago, Nycticehus, and Perodicticus the sublingua is broader 
than in the Lemuridse, especially at the apex, which is almost 
truncated and armed with many denticles — i. e. 9 in a Gulago 
monteAri^&ndi its attachment to the tongue in the middle line is 
set farther back so that a greater extent of its apex is free. I 
also noticed 9 denticles at the tip in a specimen of Perodicticus 
iheanus. But in both Perodicticits and Galago these denticles 
form a continuous series, whereas in an example of Nycticehus, 
with 7 denticles, the outer on each side was larger than the rest 
and separated from them (text-fig. 10, E, F, H). 

In Chiromys the sublingua is of a different type. It has a 
free lateral margin, but towards its abruptly attenuated tip it is 
closely adherent to tlie tongue. It is not denticulated, but ends 
in front in a firm, deflected hook which is the anterior termi- 
nation of a thick median keel or ridge, itself carrjdng two or 
three teeth, and traversing the whole of the underside of the 
sublingua back to the upper end of the lingual frenum. Just in 
front of the point there is on each side of the sublingua a. small, 
probably glandular pouch, like a watch-pocket, with its orifice 
looking forwards. The frenal lamellae are present and narrow; but 
I could not be sui'e of their exact foi-m. In the example of this 
genus I examined the sublingua is less cordate and the tongue 
less rounded than depicted by Owen, who, moreover, did not 
mention the small teeth on the carina or the pocket-like glands 
near the base of the sublingua (text-fig. 11, A, B). 

The sublingua of Tarsius, described and figured by Bui'meister, 
is difi'erent from that of Lemurs. It is not so well developed, is 
softer in consistency, and is defined from the tongue laterally by 
a groove. In the middle it is provided with a rod-like thickening 
which has a knob-like expansion at the apex. This rod, corre- 
sponding to the median carina of the sublingua seen in Chiromys 
and other Lemurs, was compai-ed to the lytta of the tongue of 
Canis by Burmeister. But the bifurcation of this rod and some 
other structures at the tip of the sublingua described by that 
author I was unable to detect in the single example of the tongue 
of Tarsius I examined. In this specimen the frenal lamellse 
consist of a pair of longish slender processes, each tipped with a 
few small projections. At the proximal end of these processes 
on the outer side the edge expands abruptly to form a flap with 
a lobidate margin which extends far backwards along the sides 
of the tongue. Burmeister described the frenal laniellse as the 



CHARACTERS OF THE LEMURS AXD TARSIUS. 



39 



" Unterzunge." His figure represents this structure as un 
undivided semielliptical lamina witli evenly denticulated margin. 
This does not agree with my observations (text-fig. 11, C-E). 

It has been suggested that the sublingua of the Lemurs acts as 
a tooth-brush to clean the porrect, close-set, and comb-like series 
of anterior mandibular teeth. Lemurs certainly use these teeth 
to comb their own fur and that of their companions. I have 
never seen them employ these teeth for any other purpose, and 

Text-fisfure 11. 




A. Side view of the tongue of Chiromys. 

B. Lower view of the same. 

C. Lower view of the tongue of Tarsius raised from the floor of the mouth. 

D. The same extracted with the frenal lamellifi and the fringe attached. 

E. Side view of the tongue of Tarsius, showing the comparatively slight 

differentiation of the sublingua and the well-developed frenal lamella; with 
backwardly extending lobulated fringe. 

since there is frequently a rapid movement of the tongue after 
the combing action, I do not doubt that the suggestion as to 
the function of the sublingua is correct ; and the suggestion is 
strengthened by the structure of the sublingua in Chiromys, 
where the median keel ends in a firm hook well adapted to all 
appearance for removing particles lodged between the two rodent 
incisor teeth of the lower jaw. 



40 MR. R. I. POCOCK ON THE EXTERNAL 

In connection witli the fur-combing use of the j^ectiniform 
mandibuUir teeth in the true Lemurs, it must be remembered 
that the spatulate finger-tips iind short nails of these animals 
deprive the digits of the scratching power they possess in ordinary 
n)ammals with narrow finger-tips and compaiatively long claws 
or nails. Only one digit in the Lemurs is functional as a 
scratcher, namely, the second of the hind-foot, which is short 
with small terminal pad and long semi-erect claw. Thus is 
established a most interesting correlation of characters in the 
Lemurs : — namely, the uselessness of the fingers for scratching 
the fur, the modification of the anterior mandibular teeth to sub- 
serve that end, and the difierentiation of the sublingua to keep 
these teeth clean from scqrf and hair. Now in Ikirsias the lower 
mandibular teeth are not modified to form a comb. Possibly 
they are employed for cleaning the coat, but their structure 
makes them less liable to be clogged, and at the same time less 
efficient instruments for the purpose than the corresponding teeth 
of the typical Lemurs. Possibly, perhaps probabl}^, for these 
reasons, the sublingua is less differentiated and two of the digits 
of the hind-foot, the second and third, are set apart as sciatchers 
and are capable of acting in unison, almost like the corresponding- 
united digits of the syndactylous Marsupials. 

The sublingua of Ghiromys probably serves mainly the purpose 
of keeping the gnawing-teeth free from woody fibre ; and, for 
anything I know to the contrary, it may also cleanse them of 
hair and scurf. But I am not aware whether these teeth are 
used as a comb or not. At all events, in the case of this genus it 
is interesting to note that the absence of the typical lem urine 
dental comb is accompanied by well-developed claws and by fur 
of a very different texture from that of ordinary Lemurs, since 
it consists of a light underwool covered by long coarse hairs. 
Possibly the usefulness of the claws in combing this fur has been 
one of the principal guiding factors in their evolution from nails. 

The Anus and its Glands. 

In all the Lemuroids examined, with one exception, the anus 
is situated in the noimal , position below the joint of the tail so 
that the basei of that organ, when depressed, closes over it. But 
in Chirogaleus majo7- it is placed below, or on the distal side of 
the joint of the tail, so that when that organ is raised the anus 
is carried up on its base (text-fig. 16, A). 

Normally in this group, that is to say in Nycticebus, Perodic- 
ticus, Galago, Ghiromys, the Indrisidfe apparent]}', and in Lemur 
catta, variegatus, and coronatus, the perineal and circumanal areas 
are covered with hair ; but in Z. macaco, L. albifrons, and the 
various forms such as nigrifrons, rufus, ftdvus, ci'iiereiceps, asso- 
ciated with albi/rons, these regions are covered in both sexes 
with nearly naked, folded, wrinkled, and glandular skin, and 
similar naked skin extends for an inch or so along the xoot of 



CHARACTERS OF THE LEMURS AND TARSIUS. 



41 



the tail (text-tigs. 12, A ; 15, E). In other species referred to 
this genus, i. e. L. mougoz and L. rubriventer, tUe glandular area 
is much less differentiate.!, the skin being n.ore closely covered 
with tine, short hair and less distinctly folded and wrinkled. 

Text-figure 12. 




A Anal and genital area of male Lemur albifrons, sliowing the nearly nated 

-Glandular skin stretching from the root of the tail to the scrotum. 
B. The ^2.meoi Lemurvwngoz, showing the hairy but partially wrinkled skin 

round the anus. • i , ^ i • 

C The same of female Leniiw mongoz, showing the absence of wrinkled skin 
■ round the anus, the long grooved clitoris (c?.), and the orifice of the urethra 
(ti.) below that of the vagina {v.). 
D. The same of male Lemur- catta, showing the hairy unwrinkled skin round the 
anus and the naked scrotum. 



42 MR. R. I. POCOCK ON THE EXTERNAL 

The only example of L. ruhriventer examined, namely a female, 
had the skin of the circumanal area distinctly bnt not strongly 
wrinkled. On the other hand, some females of L. viongoz show 
no sign of wrinkling, others show traces of it, and in some 
males the wrinkling is rather strongly pi^onounced. A complete 
gradation therefore in the development of the glandular area can 
he traced from L. coronatus through L. mongoz to L. macaco and 
L. alhifrons (text-fig. 12, A-C). 

The paired anal glands, so well known in some orders of 
Mammals, e. _r/. the Rodentia and Carnivora, appear to be absent 
in almost all Lemurs. The only genus in which I have found a 
trace of them is Chiromys, where they are represented by a pair 
of small, shallow invaginations, one on each side of the anal 
orifice. 

The External Genitalia of the Male. 

According to Milne-Edwards and Grandidier each of the three 
genera of Indrisidse may be characterised by the structure of the 
penis and baculum. The penis appears to be short, subcylin- 
drical, and apically truncated, and the baculum is distally 
biramous in all cases. 

In Lichanotus the epithelium is striated, and there are on each 
side of the organ near the middle of its length two strong 
recurved spines set one above the other. The baculum is greatly 
expanded at its proximal end and gradually narrows from the 
thickening to the middle of its length. From that point the two 
sides diverge gradually to the apices of the two branches, which 
themselves diverge evenly at about an angle of sixty degrees, 
each branch being rather less than one-third of the length of the 
whole bone. 

In Indris the epithelium of the penis is ii-regularly reticulated, 
and on each side of the organ there is a patch of rather small 
spines set in three irregularly vertical rows which extend also to 
the underside. The baculum differs from that of Lichanotus in 
being thicker in the middle of its length, without any proximal 
(or basal) expansion, and in having the two bi^anches much less 
divergent and a little longer, each slightly exceeding one-third 
the length of the entire bone (text-fig. 14, C). 

In Projnthecus the epithelium of the penis is grooved and beset 
with minute spicules. The baculum differs from that of Indris 
in having the branches very long, each being more than half the 
length of the entire bone, longer, that is to say, than the stalk 
instead of shorter as in Indris and Lichanotus. 

The penis of Ghirogaleus major (text-fig. 13, I, K, L) is rather 
short and broad, nearly parallel -sided, with an ovate extremity 
giving it a somewhat linguiform appearance from the lower or 
upper view. It is longitudinally grooved and closely punctured, 
the punctures possibly marking the position of minute spicules 



CHARACTERS OF THK LEMURS AND TARSIUS. 



43 



rubbed off. The orifice of the urethra is situated above the pad 
covering the tip of the baculum, a character in which the penis 

Text-fiyure 13. 




A. Penis of Galago crassicaudatus from below with the frill folded over the tip 

of the glans. 

B. Tip of the glans of the same with the frill spread. 

C. End of the penis of the same from the side with the tip protruding from the 

frill. 

D. Penis of Galago Senegal ensis from below. 

E. Tip of the glans of the same showing the absence of the frill. 

F. Penis of Nycticebus from the side. 

G. Tip of the glans of the same to compare with B and E. 
H. The same of Perodictimis. 

I. Penis of Chirogaleus major from below. 
K. Tip of penis of tbe same showing the orifice of the urethra above the tip of the 

baculum. 
L. Penis of the same from the side with a probe passed dowa the urethra. 
M. End of penis of Lemur macaco from below. 
N. The same from above. 
O. Penis of Lenmr alhifrons from the side. 
P. Tip of penis of the same. 



44 MR. R. I. POCOCK ON THE EXTERNAL 

differs from that of all the Leinnroid Primates examined by me. 
Above the orifice there is a transvei'se glandular depi'ession 
overlapped by a thick flap of epithelium. A somewhat similar 
crescentic flap half encircles the bacular pad laterally and below. 
The baculum (text-fig. 14, A, B) itself is a little longer than that 
of an adult example of Lemur albifi'ons more than twice the size 
of Chirogaleus onajor. Seen from the side the baculum shows a 
slight sinuous curvature, the distal half being depressed. Seen 
from above or below, the main part of the shaft is straight and 
subcylindrical, but its apical fourth is divided into two curved 
branches, a right and left, which diverge at first, then converge 
so as almost to meet apically in the middle line, circumscribing 
an ovate space. The urethra, which runs along the underside of 
the main portion of the baculum, passes through this space at 
the point of bifurcation of the baculum. Hence it comes about 
that the orifice of the urethra lies above the pad which covers the 
juxtaposed tips of the two branches. 

In being distally biramous the baculum of Chirogaleus resembles 
that of the Indrisine Lemux's, a point of great interest. It is 
most like that of Lichanotus, but has the base less expanded and 
the branches curved and convergent apically instead of widely 
divaricated. According to Beddard Hapalemxir also has a 
bifid baculum. I do not know the course of the urethra with 
regard to the baculum in the Indrisidse, nor is Beddard quite 
clear on this point in his description of the penis of Hapcdemur, 
but since he states that the urethra opens at the posterior end 
of a groove marking on the glans penis the forking of the 
baculum, it may be inferred that the orifice of the urethra is 
beneath the tip of the baculum as in typical Lemurs and not 
above it as in Chirogaleus. 

On one of the plates of his vinfinished work Milne-Edwards 
illustrates the male genitalia of a Lemur which, although un- 
named in the legend, is clearly shown to be L. catta by the naked 
scrotum. Four bacula are figured on the same plate. They 
may have been taken from specimens representing four distinct 
species or from four examples of L. catta. If the latter, the 
figures attest a certain amount of individual variation in this 
bone, particvilarly in width at the base and expansion and 
curvature at the tip. There the matter must rest. 

In the species I have examined, namely L. catta, 'macaco, 
alhifrons, and coronata, there is very little variation in the penis 
and baculum. The penis (text-fig. 13, M-P) is subcylindrical 
and armed in the middle of its length with many reversed spines 
which are mostly of small size ; two or more pairs, however, on 
each side are much larger than the others, but these large spikes 
are not always either symmetrically placed or numerically iden- 
tical on the two sides. The orifice of the urethra is terminal 
and opens just beneath the tip of the baculum. The baculum is 
a comparatively short and slender rod with a larger proximal and 



CHARACTERS OF THE LEMURS AND TARSIUS. 



45 



a smaller, sometimes incipiently bilobed distal expansion. As 
compared with the size of the animals, the baculum in the 
species referred to Lemur is smaller than in any other genns 
(text-fig. U, H). 

In Galago senegalensis the penis is long (text-fig. 13, D, E). 
Its narrow and cylindrical, proximal portion gradually expands 
distally to a considerable extent, and its terminal portion is 
attenuated. The orifice of the urethra, opening beneath and 
behind the hardened rounded pad covering the tip of the baculum, 
is provided with a small inferior and two small lateral labia. 
Except at the distal and proximal ends the epithelium of the 
penis is thickly beset with comparatively coarse reversed spines 

Text-figui^e 14. 




A. Haculum of Clnrogaleus major from the side, the line indicating' the course of 

the urethra. 

B. The same from above showing the apical bifurcation. 

C. Baculum of Indrin from above (copied from Milne- Edwards). 

D. The same of G-alagn senec/alensis from the side. 

E. The same of Galago crassieaudatus. 

F. The same of Ni/cticebns. 

G. The same of Perodicticus. 

H. The same of Lemur alhifrom^. 
I. The same of Chirnmt/s. 

(All the figures twice nat. si/e.) 



ap]3roximately equal in size. The baculum (text-fig. 14, D) is 
long and slender, being actually nearly twice as long as that of 
an adult male Lemur alhifrons. It is practically sti'aight and 
gradually attenuated fi'om its broad base, but there is a slight 
sinuosity in its distal third and a slight and gradual thickening 
at the apex which is blunt. 



46 MR. R. I. POCOCK OX THE EXTERNAL 

In Galago crassicatidattts and monteiri (text-fig. 13, A-C) the 
penis differs considerably from that of G. senegalensis. It is 
clavate in form, being grachudly incrassate from the base to the 
bhnited tip. The orifice is just below the ;ipex of the baculum, 
and the two are encircled by a frill of grooved, wrinkled epithe- 
lium, forming a sort of secondary prepuce, which is attached by a 
frenum to the lower lip of the orifice and encloses a glandular 
space. The spines covering the penis ai-e bidentate or tridentate, 
and much smaller than in G. senegalensis. The baculum (text- 
fig. 14, E), although actually longer than in G. senegalensis, is 
approximately the same shape and relatively about the same size. 
In Nycticebus the penis (text-fig. 13, F, G) is much shorter than 
in Galago and smooth and of tolei-ably even thickness through- 
out. The callous pad at the tip of the baculum is large, rounded 
and prominent, and overhangs the orifice of the urethra, which 
lies in a glandvdar space bounded laterally and below by a pair 
of labia practically as in G. senegalensis. The baculum (text- 
fig. 14, F) is a nearly straight rod, only about two-thirds the 
length of that of G. senegalensis, although the animal itself is 
much bigger. 

In Perodicticus the penis (text-fig. 13, H) is short and smooth 
and like that of Nycticebus except that the tip of the baculum 
and the orifice of the urethra are surrounded by a complete hood 
or frill of wrinkled epithelium enclosing a circular glandular 
space. The baculum (text-fig. 14, G) in the specimen examined 
is shorter than that of the example of Nycticebus, but decidedly 
thicker at the base inferiorly and with its upper edge a little 
more sinuous. 

According to Huxley s description (P. Z. S. 1864, p. 334) the 
extremity of the glans penis in Arctocebtts closely resembles that 
of Perodicticus except that the encircling hood is bifid in the 
middle line below. He states, however, that the baculum is "75 
of an inch in length, about twice as long, that is to say, as in my 
example of Perodicticus, although, judging from the dimensions 
of the limbs, his animal was considerably the smaller of the two. 
In Chiromys the penis is rugulose, wrinkled, and very slightly 
narrowed distally for three-fourths of its length, then somewhat 
abruptly attenuated to the apex, where the upturned tip of the 
baculum terminates; but just below this point there is a little 
soft, curved, subcylindrical process upon which it seems probable 
the orifice of the urethra, opens. When the prepuce is reflected 
to its fullest extent, the basal fourth of the exposed portion of the 
penis is seen to be provided with five probably glandular, longi- 
tudinal grooves, one in the middle line above and two on each 
side, one above the other, the ventral middle line being occupied 
by the frenum. The baculum (text-fig. 14, 1 ) is tolerably stout and 
longer than in any genus of Lemuroid known to me. Allowing for 
its curvature, it is as long relatively as in Galago crassicaudatus. 
When seen from the side it is markedly sinuous, its upper edge 



CHARACTERS OF THE LEMURS AND TARSIUS. ' 47 

being concave close to the base, then convex, then strongly con- 
cave owing to the iipcurling of the distal fourth of its length. 
The ventral side is curved in correspondence. No other 
Lemuroid known to nie has a strongly upcurled tip to this bone. 
In connection with the scrotum there is one fact to be re- 
corded. In Leynur catta this sac is alwa3's naked (text-fig. 12, D). 
In all other species of Lemuroid Primates it is clothed with hair 
normally (text-figs. 12, A, B; 16, A). Occasionally the postero- 
inferior portion is ]iaked, as I have noticed in one or two specimens 
of Galago ; but I have no doubt that in these cases the absence 
of the hair Avas due to rubbinff. 



The Eccternal Genitalia of the Female. 

In the standard text-books of Mammals published even as 
recently as Max Weber's in 1904, it is stated that the urethra 
traverses the clitoris in the Lemuroidea. This is not true of any 
Madagascar Lemur I have examined, and applies only to the 
Asiatic and African forms. In the Madagascar species the 
urinary orifice opens at a varying distance between the vaginal 
apei-tnre and the apex of the clitoins, generally much closer to 
that orifice and only in one case,- Lemur catta, a little nearer the 
tip of the clitoris. 

Milne-Edwards and Grandidier have shown that the clitoris 
varies considerably in form in the three genera of Indrisidfe. 
In Lichanotus (Avahis) it is long, pendnlous, and narrow, and 
parallel-sided in its distal two-thirds, but expanding proximally 
towards the orifice of the vulva. From the orifice of the urethra, 
situated a little below the vulva, a groove extends towards the 
tip of the clitoris, and at the extreme tip of the latter is placed 
the aperture of a glandular depression. 

In Projyithecus the clitoris is much shorter and thicker, with 
a bi'oadly rounded distal end carrying a glandular orifice. The 
aperture of the urethra lies approximately midway between the 
orifice of this gland below and of the vagina above. In Indris 
the distal end of the clitoris is gradually a.nd widely expanded 
laterally and extended considerably beyond the orifice of the 
gland, but the orifice of the urethra is situated even nearer to 
the vulva than in Lichanotus. 

In the species usually referred to the genus Lemur there is 
considerable variation in the structure of the genital area of the 
female. 

In L^emur varius the clitoris is a short, tliick, fleshy excrescence 
with a blunt apex and somewhat cordate in shape, rising from 
the centre of a tolerably large area of naked skin. Its free 
posterior surface is mai-ked by a median groove oi- rima defined 
laterally by a pair of thick labia. When the latter are separated 
the urinai-y channel, a gutter with thin elevated margins, is 
displayed. In a mated female this gutter is seen to lead from 



48 



MR. R. I. POCOCK ON THE EXTERNAL 

Text-fifTure 15. 




A. Anal and genital area of female Lemuv catta, showing the thick clitoris 

protruclina; between the labia of the vulva. 

B. The same of female Lemur variegatus, showing the short cordate vulva arising 

from a naked area of skin. 

C. Vulva of the same enlarged, showing the lateral grooves and the orifices of the 

urethra and vagina covered by membrane (Innnen) in the virgin female. 

D. Side view of the same vulva. 

E. Anal and genital area of female Lemur alhifrons, showing the circumanal 

glands and tlie long attenuated clitoris. 

F. The stiniu of Lemur coronatnit, showing the long clitoris and the absence of 

circumanal glands. 

G. Vulva of the same on larger scale, showing the supplementary labia beneath 

the orifice of the vagina. 

cJ., clitoris : u., orifice of urethra : v., oi'ifice of vagina. 



CHARACTERS OF THE LEMURS AND TARSIUS. 



49 



the orifice of the urethra, which is just below that of the vulva ; 
bvat in a virgin female the two orifices are concealed by a flap of 
membrane (Jiymen), below the edge of which the open channel 
extends, narrowing in its course to the apex of the clitoris. The 
sides of the clitoris are marked by a couple of deep, probably 
glandular, grooves (text-fig. 15, B, C, D), 

III Lemur catta the clitoris is a thick elongated excrescence 

Text-figure 16. 




A. Anal and genital area of male CliirngaJeus major, showing the position of the 

anus on the root of the tail. 

B. The same of female Semigalago demidoffi, showing the long pendulous clitoris 

below the vulva. 

C. The same of female Grolago crassicaudatus with the labia of the vulva spread 

open. 

D. Vulva of the same with the labia closed. 

E. Vulva of the same from the side. 

F. Anal and genital area of female Tavsius, showing the labia of the vulva 

partially spread open. 

G. The same from the side with the labia closed. 

ch, clitoris; I., labium of vulva ; ?(., orifice of urethra ; v., orifice of vagina. 

Proc. Zool. See— 1918, No. IV. 4 



50 Ml{. U. I. POCOCK ox THE EXTIillXAL 

with the orifice of the vulva at its base and that of the urethra 
about one-third of the distance from the tip — much lower, that 
is to say, than in L. varius. The urinary channel runs from the 
urethral orifice to the apex of the clitoris. The orifice of the 
vulva is bounded on each side by a pair of thick labia, which, 
when separated, diverge from above that orifice and inferiorly 
disappear in the area of naked integument from which the 
clitoris rises (text-fig. 15, A). 

In L. macaco, alhifrons, fidvus, mongoz, rufiventer, and coro- 
natits, the clitoris is narrow, elongated, attenuated towards the 
apex, and arises from an area of naked or nearly naked skin. It 
is channelled to the apex from the orifice of the urethra, which 
opens a short distance below that of the vagina. It varies to m. 
certain extent in length within specific limits, e. g. L. alhifrons, 
and is rather exceptionally long, less attenuated and narrower 
at the base in L. coronatus than in the others. Moreover, in the 
single example of this species examined, two pairs of elongated 
lamina?, constituting labia, div^erged obliquely just below the 
orifice of the vagina (text-tigs. 12, C ; 15, E, F, G). 

In the Asiatic Lemaroids the clitoris differs from that of the 
Madfigascar forms in Ijeing traversed by the urethra which opens 
at its tip. 

In Galago crass ic(tiidat as the clitoris is elongated, thick from 
before backwards in profile view, and parallel-sided and com- 
pressed from the posterior aspect : at its tip there is a depression, 
probably glandular, almost encircling the urethral orifice, and 
comparable to the similarly situated gland in the Indrisidse. 
The vulva consists of two turgid labia, of which the opposed 
surfaces are provided with several small fine laminae, running 
from the edges of the labia inwards towards the orifice of the 
vagina (text-fig. 16, C, D, E). 

In Ilemigalago the clitoris is relatively much longer and 
thinner than in G. crassicaudata. The distal portion appears to 
be strengthened by a baculum and the tip is pi'ovided with a 
glandular depression encircling the end of the bone. The two 
labia at the sides of the orifice of the vagina are relatively 
smaller than in G. crassicaudatus (text-fig. 16, B). 

In Nycticehus the clitoris is short and thick, resembling the 
prepuce of a penis. There is a glandular depression at the tip. 
The orifice of the vagina is a wide transverse rima just at the 
base of the clitoris. The general appearance of the external 
genitalia is very different from that of Galago and Hemigalago. 
Possibly the specimen examined was a virgin female. 

The female external genitalia of Tarsias are unlike those of 
the Asiatic and Masearene Lemurs, but in the single example 
examined recall rather those of the Old World pithecoid Primates. 
The vulva is a laterally compressed, elongated excrescence, with- 
out pendulous apex. The rima is short and bounded by a pair 
of labia concealinsf the small clitoris and the orifices of the 



CHARACTERS OF THE LEMURS AND TARSIU6, 01 

Tirethra and vagina, which are comparatively close together. It 
is significant that the vulva of this genus as a whole is less like 
the vulva of the Galagos thftn that of the true Lemurs. It 
■approaches most nearly the vulva of L. varms (text-fig. 16, F, G). 



General Conclusions. 

The conclusions of general interest which have suggested them- 
selves in the course of the investigations above detailed are 
subjoined in the order of their importance : — 

1 . In the case of Tarsius the structure of the upper lip and of 
■the nose severs the genus completely from the Lemurs and brings 
it into line with the Pithecoid Primates. Taking this character 
in conjunction with the nature of the placenta, the presence of 
ihe postorbital partition, and other well-known features, it seems 
that Hubi-echt was quite right in removing Tarsius from the 
Lemurs and placing it in the higher grade of Primates. I pro- 
pose to give practical expression to this view by dividing the 
Primates into two great series. For the first, comprising the 
Lemurs, the old name Strepsirhini is available. For the second, 
■comprising Tarsius and the Pithecoidea, I suggest the title 
Haplorhini. The Haplorhini will contain two divisions, the 
Tarsioidea and the Pithecoidea*. 

2. With Tarsius eliminated from the Lemurs, the Strepsirhini 
may be divided into the Chiromyoidea and the Lemuroiclea. It 
has been the fashion of late years to depreciate the characters of 
Chiromys. Dr. Standing indeed gave the genus merely subfamily 
rank under the Indrisida?, the latter being equivalent to the 
Lemuridse. But that classification sacrificed the characters of 
Chiromys to the hypothesis that the genus is a specialised off- 
shoot of the Indrisoid stock. That may be true. Nevertheless, 
the specialisation has proceeded so far, and in so many directions, 
that it appears to me impossible to dispute the claim that the 
Lemuridse and the Indrisidpe are much more nearly akin to each 
-other than either is to the Chiromyidfe. In the description of 
Chiromys the peculiarities of the teeth generally distract attention 
from the curious cranial modifications correlated with the rodent 
■dentition, such as the immense size of the premaxillse, which 
reach the lachrymals and exclude the reduced maxilla; from con- 
tact with the nasals ; also the absence of the bony ridge closing 
the glenoid behind and the longitudinal extension of the man- 
dibular condyle, two correlated characters subservient to the 
back and forth movement of the mandible well known in the 

* I {ji-efer tliis name to Anthropoidea because in ordinary terminolog}' the title 
" antliropoid," reasonably according; to its meaning, has become restricted to the 
man-like Apes. A marmozet can hardly be called " anthropoid," with any approach 
to the real meaning of the word. But a marmozet and a man are alike "pithecoid." 



52 Mil. R. I. POCOCK ON THE EXTERNAL 

Rodentia. These cranial features distinguish Chiromys from all 
Lemurs ; and when taken in conjunction with the teeth, with the 
peculiarities of the hands and feet, and of the sublingua, they out- 
weigh, in my opinion, the known differences between the true 
Mascarene Lemurs (Lemurida? and Indrisidse) and the Asiatic 
and African Galagos, Pottos, and Lorises. 

3. With regard to the Galagos, Pottos, and Lorises, I am oidy 
acquainted with one invariable cranial chai"acter distinguishing 
them from the Lemuridje and Indrisidae. This was pointed out 
by Forsyth Major and has been briefly expressed by Mr, Gregory* 
as follows : — In the Asiatic forms the ectotympanic is enlarged 
and external to the bulla of which it forms the outer wall. In 
the Mascarene forms the ectotympanic is inclosed within the- 
bulla, forming a ring or horseshoe. To this difference may be 
added the one pointed out above in connection with the clitoris, 
Avhich in the Asiatic genera is traversed by the urethra, whereas 
in the Mascarene forms the urethra opens above the tip of the 
clitoris. 

In view of these facts, I should divide the Lemuroidea into 
two series, for which Mr. Gregory's names Lemuriformes for the 
Lemuridie and Indrisidse, and Lorisiformes for the Loiisidse (or 
Nycticebid?e) and the Galagidse may be adopted. Similarly 
for the subdivision of the Lemuridfe I follow Mr. Gregory in 
relegating the genera to two subfamilies, the Lemurinfe and 
Chirogaleinse ; but I cannot agree with him that Hapalemiir 
belongs to the ChivogaleinBe. That genus appears to me to be 
essentially a Lemurine, its inclusion in the Chirogaleinee spoiling 
the definition of the subfamily. 

4. As a matter of minor interest it is quite clear that the- 
genus Lemur as generally admitted and as recognised in this 
paper is susceptible of division into two or three genera. L. ccdta, 
for instance, differs from the other species in having the glands 
on the fore-limb and the naked heel and scrotum, and also 
in the structure of the vulva. L. variegatus is also peculiar in 
the structure of the vulva. Furthermore, -the Galagos of the 
G. senec/alensis-ty-pe may be distinguished by the structure of the 
penis from that of the c)-assicai{,dcUus-tjY)e. Generic names 
appear to be available for these subdivisions of Lemur and 
Galago ; but I do not propose to enter into that question now. 

My views above set forth differ in so many particulars from 
those of Mr. Gregory that it may be interesting to tabulate our 
classifications side by side, omitting those he adopts based upon 
extinct genera, which, so far as I am aware, do not materially 
affect the arrangement of recent forms. 

* Bull. Geol. Soc. Amev. 26, pp. 432-436, 1915. See also Bull. Aniev. Mas. Nat. 
Hi-st. 35, pp. 266-267, 1916. 



CHARACTERS OF THE LEMURS AND TARSIUS. 



53 



Gregory, 1915-1916. 



^Ordev PRIMATES. 

Subord. Lemueoidea. 
Series LEMUEiFOfiMEri. 
Fam. Lemurid^. 
Subfaui. Lemiirince. 
), Chirogalehia. 
Fam. Indkisid^. 
„ Chieomtid^. 
■Series Loeisifoemes. 
Fam. LoRiSEDiE. 
Subfam. Lorisince. 
„ GalagiiKS. 

Series Taesiifoemes. 
Fain. Taesiid^. 
'Subord. Antheopoidea. 



Pocock, 1918. 



Order PRIMATES. 
, Grade Steepsiehini. 
Subord. Lemuro'idea. 
Series Lemueifoemes. 
Fam. Lemueidji. 
Subfam. Lemurinee. 
„ ChiroffaleitKe. 
Fam. Indkisid^. 
Series Loeisifoemes. 
Fam. LoEisiD^. 
„ Galagidj?. 
Subord. Chieomtoidea. 
Fam. Chieomtidj?. 
Grade Haploehini. 
Subord. Taesioibea. 

PiXHECOIDEA. 



Apart from the removal of Tarsius from the Lemuroidea ami 
the union of the Lorisidfe and Galagidae in a group equivalent to 
the Lemuridfe and Indrisidse combined, my classification agrees 
in the main with the classification published in Flower and 
Lydekker's ' Mammals,' and in other works of about that period. 
Of more recent classifications it appears to me that Gadow's 
.(Class, of the Vertebrata, 1898, pp. 52-53), although brief, ex- 
presses the facts with the nearest approach to the truth, especially 
in the primary division of the Primates into the three suborders 
Lemures, Tarsii, and Simile. 



0\ THE PYRALID.E, SUBFAMILY HYPSOTKOPIN^. 



5. A Classification of the Ptralid^, subfamily Hypso- 
TROPiN.E. By Sir George Hampson_, Bart., F.Z.S. 

[Received Februaiy 19, 1918 : Read March 5, 1918.] 



Proboscis aborted or absent ; palpi upturned, oblique or down- 
curved, the males of the species with the maxillary palpi brush- 
like and contained in a hollow of the labial palpi usually having 
the palpi oblique in the male and downcurved in the female ; 
maxillary palpi small and filiform, well developed and more or 
•less dilated with scales at extremity, or brush-like and contained 
in a hollow of the labial palpi ; frons smooth, with tuft of hair, 
or prominences of various forms ; antennae of male ciliated, 
laminate, serrate or pectinate with uniseriate bi-anches, the 
basal joint often dilated and the shaft often downcurved at base 
with a ridge of scales in its sinus; the build slender; tibise with 
all the spurs present ; abdomen smoothly scaled. Fore wing 
narrow ; vein 1 a separate from 16; 1 c absent ; veins 2, 3 
rarely stalked ; vein 4 often absent or stalked with 5, 3 and 5 
stalked or from the cell; 6 from below upper angle; 7 absent; 
8, 9 and often 10 stalked or 9 absent, rarely 10 also absent, 
8 given off before or after 10; 11 from cell. Hind wing with 
the median nervure pectinated on upper side ; veins 1 a, b, c 
present; 2 from near or well before angle of cell; veins 3 and 4 
sometimes absent, or 3 and 5 stalked or from the cell, 4 often 
absent, or 3, 4, 5 all present, 4 stalked with 5 or from cell ; 6, 7 
stalked or from upper angle of cell ; 8 anastomosing with 7 or 
approximated to it but free. 

The larvte of very few species are known; these live on 
Graminacece and pupate in the ground in a chamber formed 
by agglutinated particles of the soil ; Einmalocera depressella, 
however, is injurious to sugar-cane, the larva feeding in its 
roots. 

The su1)family is a development from the Anerasticmce — Ane- 
rastia Hiibn. Yerz. p. 367 (1827), type dignella Hiibn. — from 
which it differs only in the proboscis being absent or aborted 
and non-functional. 

The types of the new species are in the British Museum, the 
species of which the types are in the collection are mai'ked with 
a t, and those not in the collection with a *. 

No quotations from German authois published since Aug. 1st,. 
1914 are inserted. " Hostes humani generis.'''' 



56 



SIR GEORGE HAMPSON ON THE 



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PYRALID^, SUBFAMILY HYPSOTROPIN^. 



Key to the Genera. 



57 



A. Hind wing with veins 3 and 4 absent. 

a. Fove wing with vein 8 fi'om 9 before 10. 3 and 5 stalked Coenochroa, 

b. Fore wing with vein 8 from 9 after 10. [p- 58. 
«! Fore wing wdth veins 3 and 5 stalked. 

k Palpi npturned Mennthia, yi.m. 

b- Palpi oblique, straight Cajem, p. o9. 

c2. Palpi downcurved Statma, ^. 5^. 

Ji Fore wing with veins 3 and o from the cell. 

a2. Palpi upturned Tmfrasifra, p. 61. 

b'. Palpi porrect. . , o 

«:< Fore wing with vein 9 absent, 11 stalked with 8 

a,,jl 10 Coenot7-opa, \).S9. 

b^ Fore wing with vein 9 present, 11 from cell Mesodiphlehia, 

LP- «1- 
c Fore wing with vein 10 from the cell Calamotropa, 

Lp- "■^• 

B. Hind wing with vein 3 present, 4 absent. 
a. Fore wing with vein 4 absent. 

rti Fore wing with veins 9 and 10 absent, 8 from cell, 3 and 

5fromcell ^ ra7»_pa, p. 62. 

61. Fore wing with vein 9 absent, 10 from cell, 3 and 5 

from cell Bandera, p. 89. 

ci. Fore wing witli vein 8 from 9 before 10, 3 and 5 stalked... Hosidia, p. 63. 
cJi. Fore wing with vein 8 from 9 after 10. 
a'. Fore wing with veins 3 and 5 stalked. 
flS. Palpi upturned. . . 

a-*. Fvons with rounded prominence Dewfeea, p. b4. 

b*. Frons without prominence. 

a''. Palpi reaching to above vertex of head, the 2nd 

joint slenderly scaled Ceara, p. 63. 

65. Palpi not reaching to above vertex of head, the 

2nd joint broadly scaled Leotropa, p. 64. 

6^. Palpi oblique or downcurved. 
n-*. Fr6ns with corneous plate produced to two slight 

points in front Alamosa, Y>- b5. 

b-i. Frons without corneous plate JVa vasuta, p. 6o. 

6-. Fore wing with veins 3 and 5 from cell. 

a3. Hind wing with veins 3 and 5 stalked Kypsotropa, 

63. Hind wing with veins 3 and 5 from the cell. [p. 67. 

a*. Palpi upturned Sudania,^. m. 

l\ Palpi downcurved Hhodoclirysa 

<-y. Fore wing with vein 10 from the cell, 8 and 9 stalked. [P- ^'^^ 

a". Hind wing with veins 3 and 5 stalked. 

a3. Frons with flattened corneous plate, rather trifid in 

front and produced to short lateral points, a cor- LP- "°- 

neous plate below the frons BapJiimetopus, 

6^. Frons with pointed conical prominence Chortonceca, 

c^. Frons with long truncate corneous process with raised LP- 80. 

rim at extremity 3Ief.acrateria,_ 

d^. Frons with rounded prominence with raised rim at Lp- '^• 

extremity Prinanerastia, 

[p. 80. 

e3. Frons without proininence Bhinaphe, p. 82. 

b'. Hind wing with veins 3 and 5 from the cell. 

a3. Hind wing with vein 8 anastomosing with 7 Lanrentia, p. ,90. 

63. Hind wing with vein 8 not anastomosing with 7 Epidatiria,v-9l. 

■i. Fore wing with vein 4 present. 

a*. Fore wing with vein 10 stalked with 8. 

a". Fore wing with veins 4, 5 stalked or from a point. 
a3. Hind wing with vein 2 from or from close to angle 

a"*. Frons with conical prominence Fossijrontia, 

V. Frons without prominence. '-^ine 

aK Frons with long pointed tuft of hair Aurora, ^^.Wb. 

bK Frons without tuft of hair Commotrm, 

[p. 107. 



58 SIR OEORGK IIAMPSOX ON THE 

b'K Hind wing: with vein 2 from well l)efore angle of cell. 

tc*. Froiis with rounded prominence Siboffa, p. 112. 

b*. Frous without prominence, 
a"". Fore wing with veins 4, 5, hind wing with veins 

3 and 5 stalked Emaflieudes, 

h^. Fore winn- with veins 4, 5, hind wing: with veins [p. 114. 

3 and 5 from angle of cell Baptotropa, 

b-. Fore wing with veins 4, o separate. [p. 116. 

«•*. Hind wing with vein 2 from near angle of cell Biafra, p. 115. 

b'-^. Hind wing with vein 2 from well before angle of cell. [p. 116. 

a^. Hind wing with vein 8 anastomosing with 7 TSthiotropa, 

b-^. Hind wing with vein 8 not anastomosing with 7 ... Patna, p. 117. 
S'. Fore wing with vein 10 from the cell. 
a-. Fore wing with veins 4, 5 stalked. 

a^. Hind wing with veins 3 and 5 stalked Saluria, -p.d'S. 

h^. Hind wing with veins 3 and 5 from the cell ProjjJitasia, 

b". Fore wing- with veins 4, 5 from the cell. [p. 104. 

a^. Hind wing with veins 3 and 5 stalked ; frons with 

truncate conical prominence lleffalopJiofa, 

b^. Hind wing with veins 3 and 5 from the cell. [p. 117. 

a"*. Frons with prominence hollowed out in front Martia, p. 118. 

ft-*. Frons with disk of scales Discofrontia, 

[p. 118. 

c^. Frons without disk of scales or prominence Critonia,'p. 119. 

C. Hind wing with vein 4 present. 

a. Fore wing with vein 10 stalked with 8. [p. 121. 

fli. Hind wing with vein 8 anastomosing witli 7 Monoctenocera, 

J'. Hind wing with vein 8 not anastomosing with 7 Saborma, p. 122. 

b. Fore wing w-ith vein 10 from the cell. 

a'. Fore wing with veins 2, 3 stalked OsrtCi«, p. 123. 

S'. Fore wing with veins 2, 3 from the cell. 

a-. Frons with small rounded prominence with a corneous 

plate below it Ragonotia, 

b^. Frons without prominence. [p. 123. 

«(■*. Fore wing with vein.'* 4, 5 stalked Po7i/oc7ia, p. 124. 

b*. Fore wing with veins 4, 5 from the cell JSmmalocera, 

[p. 126. 

Genus CCEN'OCHROA. 

Type. 

Coenochroa Rag. N. Am. Phyc. p. 20 (1887) ccdiforniella^ 

Petalv.ma Hulst, Trans. Am. Ent. Soc. xvii. 

p. 215 (1890) illihella. 

Proboscis absent ; palpi downcurved, about three times length 
of head and thickly scaled ; maxillary palpi minute and filiform ; 
frons with conical prominence, thickly scaled above ; antennaj 
of male laminate and almost simple. Fore wing narrow and 
elongate, the apex rounded ; vein 2 from close to angle of cell ; 
3 and 5 strongly stalked, 4 absent ; 6 from well below upper 
angle; 8, 9, 10 stalked, 8 from before 10; 11 from cell. Hind 
wing with vein 2 from close to angle of cell ; 3 and 4 absent, 
5 sometimes not reaching the termen ; 6, 7 from upper angle ; 
8 stronglj' anastomosing with 7. 

(1) *CCENOCHROA ILLIBELLA. 

Anerastia illihella Hulst, Ent. Am. iii. p. 138 (Oct. 1887); 
B,ag. Rom. Mem. viii. p. 419, pi. 45. f. 9; Dyar, Cat. Lep. IST. 
Am. p. 440. 

Coenochroa -pur icostella Rag. N. Am. Phyc. p. 20 (Dec. 1887). 

U.S.A., Texas, Arizona. 



PYRALID.E, SUBFAMILY HYPSOTKOriN^:. 59 

(2) CCE^^OCHROA INSPERGELLA. 

Ccenochroa inspergella Rag. N. Am. Phyc. p. 20 (1887);. id. 
Rom. Mem. viii. p. 419, pi. "45. f. 10; Dyar, Cat. Lep. N. Am. 
p. 440. 

U.S.A., Texas, Colorado, Arizona. 

(3) *CCENOCHROA CALIFORNIELLA. 

Ccenochroa californiella Rag. IST. Am. Phyc. p. 20(1887); id. 
Rom. Mem. viii. p. 420, pi. 45. f. 8; Dyar, Cat. Lep. N. Am. 
p. 441. 

U.S.A., Colorado, California, Arizona ; Mexico, Presidio. 

A^lctorum. 

Ccenochroa monomacula Dyar, Pr. U.S. Nat. Mus. xlvii. p. 348 
(1913) Panama. 

Genus Calera. 

Type. 
Calera Rag. Nouv. Gen. p. 50 (1888) punctiliinbella^ 

Proboscis aborted and minute ; palpi downcurved, extending 
about twice the length of head and smoothly scaled ; maxillary 
paljsi minute and filiform ; frons smooth ; antennae of male 
somewhat laminate and ciliated, the basal joint somewhat en- 
larged and flattened on outer side. Fore wing long and narrow,, 
the apex i-ounded ; vein 2 from near angle of cell; 3 and 5 
strongly stalked, i absent ; 6 from below vipper angle; 8, 9, 10 
stalked, 8 from beyond 10; 11 from cell. Hind wing with 
vein 2 from close to angle of cell ; 3 and 4 absent ; 6, 7 stalked ; 
8 anastomosing strongly with 7. 

Calera punctilimbella. 

Calera punctilimhella Rag. Nouv. Gen, p. 50 (1888); id. Rom. 
Mom. viii. p. 417, pi. 40. f. 20 ; Dyar, Cat. Lep. N. Am. p. 441. 
U.S.A., N. Carolina, Iowa, Texas. 

Genus Statin A. 

Type. 

Statina Rag. N. Am. Phyc. p. 19 (1887) roseotinctella. 

Proboscis aborted and minute; palpi downcurved, extending 
about two and a half times length of head and rather roughly 
scaled ; maxillary palpi minute, filiform ; frons with tuft of 
scales ; antenna3 of male laminate and ciliated, the shaft thickened 
with scales above at base, the basal joint long, dilated and 
flattened. Fore wing long and narrow, the apex rounded ; 
vein 2 from near angle of cell ; 3 and 5 strongly stalked, 
4 absent ; 6 from below upper angle ; 8, 9, 10 stalked, 8 from 
beyond 10, 11 from cell. Hind wing with vein 2 from close to 



■60 SIR GEOllGE HAMP80N ON THE 

aiigle of cell ; 3 and 4 absent ; 6, 7 from upper angle ; 8 anasto- 
mosing strongly with 7. 

(1) *Statina gaudiella. 

Statina gaudiella Hulst, Trans. Am. Ent. Soc. xvii. p. 216 
(1890); Rag. Rom. Mem. viii. p. 415, pi. 51. f. ^; Dyar, Cat. 
Lep. N. Am. p. 440. 

U.S.xi., Texas. 

(2) Statina roseotinctella. 

Statina roseotinctella Rag. N. Am. Phyc. p. 19 (1887); id. 
Rom. Mem. viii. p. 416, pi. 47. f. 7 ; Dyar, Cat. Lep. N. Am. 
p. 440. 

U.S.A., Florida, Texas. 

(3) tSTATINA PUNCTILINEELLA. 

Statina punctilineella Hmpsn. Rom. Mem. viii. p. 416, pi. 52. 
f. 13(1901). 
S. Brazil. 

(4) tSTATINA BIFASCIELLA. 

Statina hifasciella Hmpsn. Rom. Mem. viii. p. 416, pi. 52. 
1 8 (1901). 
U.S.A., Texas. 

(5) Statina rhodobaphella. 

Statina rhodobaphella Rag. Nouv. Gen. p. 50 (1888) ; id. Rom. 
Mem. viii. p. 417, pi. 45. f. 6, 

Celebes, Sangir I. ; N. Guinea ; Queensland. 

(6) tSTATINA ROSINELLA, Sp. U. 

5 . Head and thorax purplish pink ; abdomen whitish suf- 
fused with I'ed-brown, dorsallj' fulvous towards base; antennaj, 
pectus, and legs whitish tinged Avith red-brown. Fore wing 
purplish pink, the veins streaked with white ; a brown shade 
below basal half of cell. Hing wing ochreous white. 

N. Nigeria, Minna {Macfie), 1 $ type. Exp. 16 mm. 

(7) tSTATINA CASHMIRALIS. 

Statina cashmiralis Hmpsn. J. Bomb. Nat. Hist. Soc. xv. 
p. 20(1901). 
Kashmir. 

Genus Menuthia. 

Type. 
Menuthia Rag. Nouv. Gen. p. 50 (1888) nanella. 

Proboscis absent ; palpi upturned to far above vertex of head, 
slender and smoothly scaled ; maxillary palpi filiform ; frons 



PYRALID^, SUBFAMILY HYPSOTKOPIN^E. 61' 

smooth ; antennae of female almost simple. Fore wing long and 
narrow, the apex rounded ; vein 2 from angle of cell ; 3 and 5 
stalked, 4 absent; 6 fi-om below upper angle; 8, 9, 10 stalked, 
.8 from beyond 10 ; 11 from cell. Hind wing with the cell short ; 
veins 2 and 5 stalked, 3 and 4 absent; 6, 7 from upper angle;. 
8 strongly anastomosing with 7. 

*MeNUTHIA NANELLA. 

Menuthia ncmella Rag. Nouv. Gen. p. 50 (1888); id. Rom. 
Mem. viii. p. 418, pi. 40. f. 22. 
Zanzibar. 

Genus TiNERAsxiA. 

Type. 

Tinerastia Hmpsn. Rom. Mem. viii. p. 414 (1901)... Jlssirella. 

Proboscis absent; palpi upturned to rather above vertex of 
head .and nearly smoothly scaled ; maxillary palpi minute and 
filiform ; frons with I'ounded prominence ; antennae of male 
laminate and ciliated, the basal joint rather long. Fore wing 
rather long and naiTOW, the apex rounded ; vein 2 from towards 
angle of cell ; 3 and 5 from angle, 4 absent ; 6 from below upper 
angle; 8, 9, 10 stalked, 8 from bej^ond 10; 11 from cell. Hind 
wing with veins 2 and 5 from angle of cell, 3 and 4 absent ; 6, 7 
stalked ; 8 anastomosing strongly with 7. 

(1) tTlNERASTIA DISCIPUNCTELLA. 

Menuthia disclpunctella Hmpsn. Moths Ind. iv. p. 52 (1896) ; 
id. Rom. Mem. viii. p. 414, pi. 52. f. 10. 

Ceylon. 

(2) tTlNERASTIA FISSIRELLA. 

Menuthia Jlssirella Hmpsn. Moths Ind. iv. i^. 52 (1896); id. 
Rom. Mem. viii. p. 414, pi. 52. f. 9. 
Ceylon. 

Genus Mesodiphlebia. 

Tj'pe. 
Mesodiphlebia Zell. Hor. Soc. Ent. Ross. xvi. 

p. 251 , crassivenia. 

Proboscis absent; palpi poiTect, about twice the length of 
head, almost straight and thickly scaled ; maxillary palpi well 
developed and slightly dilated with scales ; frons with conical 
tuft of hair; antennae of male laminate and . ciliated, the shaft 
somewhat curved at base and thickened with scales in the sinus, 
the basal joint rather elongate and flattened. Fore wing long 
and narrow, the apex rounded ; vein 2 from near angle of cell 
3 and 5 separate, 4 absent ; 8, 9, 10 stalked, 8 from beyond 10 
11 from cell. Hind wing with veins 2 and 5 fi'om angle of cell 
3 and 4 absent ; 6, 7 from upper angle ; 8 anastomosing strongly 
with 7. 



62 sir george hampson on the 

(1) *Mesodipjilebia deliquella. 

Anerastia deliquella Zell. Isis, 1848, p. 861 ; Rag. Rom. M6m. 
^iii. p. 413, pi. 45. f. 5. 

Anerastia trinotelJa Berg, An. Soc. Arg. 1885, p. 275. 
Argentixa. 

(2) tmesouiphlebia crassivenia. 

Anerastia crassivenia Zell, Hor. Soo. Ent. Ross. xvi. p. 251, 
pi. xii. f. 52 (1881); Rag. Rom. Mem. viii. p. 415, pi. 40. f. 16. 
Colombia. 

(3) Mesodiphlebia stricticostella. 

Mesodiphlebia stricticostella Rag. Ann. Soc. Ent. Fr. 1887, 
p. 260 ; id. Rom. Mem. viii. p. 413, pi. 40. f. 15. 
N. Nigeria ; Sudax. 

(4) t Mesodiphlebia rosella. 

Calera rosella Hmpsn. Moths Ind. iv. p. 53 (1896) ; id. Rom. 
Mem. viii. p. 412, pi. 52. f. 7. 
Madras, jSTilgiris. 

(5) tMESODIPHLEBIA OCHRACEELLA, SpT n, 

5 . Head and thorax brownish ochi-eous ; pectus, legs, and 
abdomen white suffused with brownish ochreous. Fore wing 
uniform brownish ochreous. Hind wing white with a faint 
ochreous tinge. 

Argentina, Corrientes, Goya (Perrens), 1 $ type. Exp. 
20 mm. 

Atcctorii,7n. 

Calera alhicostella Grossbeck, Bull. Am. Mus. Nat. Hist.xxxvii. 
p. 134(1917) U.S.A., Florida. 

Genus Tampa. 

Type. 
Tampa Rag. N. Am. Phyc. p. 19 (1887) dimediatella. 

Proboscis aborted and small ; palpi obliquely upturned to well 
above vertex of head and thickly scaled ; maxillary palj)i well 
developed and dilated with scales; frons smooth; antennse of 
male ciliated. Fore wing long and nai'row ; the costa highly 
arched, the apex rounded, the termen very oblique ; vein 2 from 
close to angle of cell ; 3 and 5 separate, 4 absent ; 6 from below 
upper angle; 9 and 10 absent ; 11 from close to 8. Hind wing 
with vein 2 from near angle of cell ; 3 and 5 strongly stalked, 
4 absent ; 6, 7 from upper angle ; 8 anastomosing with 7 to 
near apex. 



pyralid.e, subfamily hypsoteopin^. 63 

Tampa dimedutella. 

Tampa dhnediatella Rag. N. Am. Phyc. p. 20 (1887) ; id. Rom. 
Mem. viii. p. 411, pi. 45. f. 4 ; Dyar, Cat. Lap. N. Am. p. 440. 

U.S.A., Florida, Texas, 

Genus Ceara. 

Type. 
Ceara Rag. Nouv. Gei) . p. 45 ( ] 888) discinotella. 

Proboscia absent ; palpi upturned to rather above vertex of 
bead and moderately scaled ; maxillary palpi small and filiform ; 
frons smooth, with large conical tuft of scales ; antennae of male 
laminate and ciliated, the shaft with small tuft of scales above 
at base. Fore wing narrow, the apex rounded : A^ein 2 from 
angle of cell ; 3 and 5 stalked, 4 absent ; 6 from below upper 
angle; 8, 9, 10 stalked, 8 from beyond 10; 11 from cell. Hind 
wing with vein 2 from near angle of cell ; 3 and 5 strongly 
stalked, 4 absent ; 6, 7 stalked ; 8 strongly anastomosing with 7. 

(1) *Ceaiia discinotella. 

Ceara discinotella Rag. ISTouv. Gen. p. 45 (1888) ; id. Rom. 
Mem. viii. p. 368, pi. 39." f. 3. 
S. Brazil. 

(2) tCEARA ALBIFASCIATA, Sp. 11. 

S . Head white ; thorax white tinged with reddish brown ; 
abdomen white tinged with brown and dorsallj^ with fulvous 
yellow towards base : pectus and legs white tinged with brown. 
Fore wing with white costal fascia narrowing to a point before 
apex and with vein 12 defined above and below by grey-brown, 
the area below it to just below the cell and vein 2 white sufi\ised 
with grey-brown, leaving the veins streaked with white; the 
inner area pale pink with a white streak on vein 1. Hind 
wing white, tinged with brown except on inner area. 

Br. E. Africa, Eb Urru (Bettou), 1 S type. Exp, 26 mm. 

Genus HosiDiA. 

Type. 
Hosidia Hmpsn. Rom. Mem. viii. p. 408 (1901). ochrineur-ella. 

Proboscis absent; palpi downcurved, extending about one and 
a half times length of head and roughly scaled ; maxillary palpi 
minute and filiform ; frons smooth, with conical tuft of hair ; 
antennae of male ciliated, the shaft with sinus and tuft of scales 
at base. Fore wing narrow, the apex rounded ; vein 2 from 
close to angle of cell ; 3 and 4 stalked, 5 absent ; 6 from below 
upper angle; 8, 9, 10 stalked, 8 from before 10; 11 from cell. 
Hind wing with vein 2 from close to angle of cell ; 3 and 5 
strongl}^ stalked, 4 absent ; 6, 7 stalked : 8 anastomosing strongly 
with 7. 



64 SIR GEORGE HAMPSON ON THE 

*H0SimA OOHRINEURELLA. 

Hosidia ochrineurella Hmpsii. Koni. Mem. viii. p. 409, pi, 45,. 
f. 1 (1901). 
Natal. 

Genus Dembea. 

Type. 
Dembea Rag. Nouv. Gen. p. 45 (1888) venulosella.^ 

Probcscis aborted and minute ; palpi upturned to about vertex 
of head and thickly scaled ; maxillary palpi minute and slightly 
dilated with scales; frons with large rounded prominence thickly 
clothed with scales ; antennae of male laminate and ciliated, the 
shaft with ridge of scales at base. Fore wing narrow, the apex 
i-ounded ; vein 2 from towai-ds angle of cell ; 3 and 5 shortly 
stalked, 4 absent; 6 from below upper angle; 8, 9, 10 stalked,. 
8 from beyond 10; 11 from cell. Hind wing with vein 2 from 
jvngle of cell ; 3 and 5 stalked, 4 absent; 6, 7 stalked ; 8 anas- 
tomosing strongly with 7. 

tDEMBEA VENULOSELLA. 

Dembea venalosella Rag. Nouv. Gen. p. 45 (1888); id. Rom.. 
Mem. viii. p. 3(58, pi. 44. f. 19. 

N. Nigeria ; Abyssinia ; Br. C. Africa. 

Genus Leotropa, nov. 

Type, L. j^hcenicias. 

Proboscis absent ; palpi obliquely upturned to about vertex of 
head and moderately scaled ; maxillary palpi minute and filiform ; 
frons smooth, rounded ; antennae of male laminate and ciliated, 
with a sinus at base containing a double ridge of scales. Fore 
wing narrow, the apex rounded ; vein 2 from towards angle of 
cell ; 3 and 5 stalked, 4 absent; 6 from below upper angle ; 8, 9, 10 
stalked, 8 from beyond 10 ; 11 from cell. Hind wing with vein 2 
from just before angle of cell ; 3 and 5 very strongly stalked,, 
4 absent; 6, 7 stalked ; 8 strongly anastomosing with 7. 

(1) tLEOTROPA phcenicias, sp. n. 

(S . Head and thorax purplish pink ; abdomen pale brownish 



brownish tinged witli purplish pinK. J^ ore wing purpiisn pinK, 
the veins streaked with white; a dark brown fascia through 
Tipper part of cell to apex, the costal area towards apex white» 
Hind wing brownish white ; a slight brown terminal line. 
Sierra Leoxe {Clements), 1 6 type. J^xp. 22 mm. 

(2) tLEOTROPA SARCINA, Sp. n. 

cJ . .Head and thorax white mixed with ochreous brown ; 
abdomen white, dorsally faintly tinged with fulvous towards 



PYRALID^, SUBFAMILY HYPSOTKOPIN^. 65 

base ; pectus and legs white suffused with brown. Fore wing 
white tinged with purplish pink, the veins streaked with white ; 
the costal area white narrowing to a point at apex and with vein 12 
defined on each side by brown, a brown shade below the costal 
area. Hind wing white with a faint ochreous brown tinge on 
costal area and in submedian interspace. > 

Br. E. Africa, Teita {Jackson), 1 S type. Exp. 16 mm. 

(3) tLEOTRDPA PAPUANENSIS, sp. n. 

(S . Head, thorax, and abdomen whitish suffused with reddish 
brown, the last dorsally tinged with fulvous towards base. Fore 
Aving whitish irrorated with reddish brown, the costal area rather 
whiter, narrowing to a point at apex and with slight brown 
streak below it. Hind wing white slightly tinged with brown, 
the cilia white. 
_ Dutch N. Guinea, Ron I. [Doherty), 1 d type. E.vj^. 14 mm. 

Genus Alamosa. 

Type. 
Alamosa Hmpsn. Rom. Mem. viii. p. 369 (1901) . injjeratella. 

Proboscis aborted and minute ; palpi downcurved, extending- 
about twice the length of head and thickly scaled ; maxillary 
palpi minute and filiform ; frons with flattened corneous plate 
produced to two slight points in front and thickly scaled above ; 
antennfe of male laminate. Fore wing narrow, the apex rounded; 
vein 2 from towards angle of cell ; 3 and 5 stalked, 4 absent ; 6 
from below upper angle ; 8, 9, 10 stalked, 8 from beyond 10 ; 
11 from cell. Hind wing with A'ein 2 from just before angle of 
cell ; 3 and 5 strongly stalked, 4 absent ; 6, 7 from upper angle ; 
8 strongly anastomosing Avith 7, 

(1) Alamosa piperatella. 

Alamosa piper atella Hmpsn, Rom. Mem. viii. p. 369, pi. 51. 
f. 25 (1901). 

U.S.A., Colorado, N'ew Mexico. 

(2) *Alamosa bipunctella. 

Alamosa bipunctella Barnes & McD. Contr. Nat. Hist. Lep. 
K Am. ii. 4, p. 184, pi. 1. f. 7 (1913). 
U.S.A., Florida. 

Genus Navasota. 

Type. 
Navasota Rag. N. Am. Phyc. p. 1 8 (1 887) hehetella. 

Proboscis aborted and miiiute ; palpi extending about the 
length of head and moderately scaled, the 2nd joint oblique, the 
3rd porrect ; maxillary palpi minute and filiform ; frons smooth 
with conical tuft of hair ; antennae of male laminate and ciliated, 

Proc. Zool. Soc— 1918, No. Y. 5 



66 SIR GEORGE HAMPSON ON THE 

the shaft with sinus at base containing a ridge of scales. Fore 
wing narrow, the apex rounded ; vein 2 from towards angle of 
cell ; 3 and 5 stalked, 4 absent ; 5 fi'om below upper angle ; 8, 9, 
10 stalked, 8 from beyond 10; 11 from cell. Hind wing with 
vein 2 from close to angle of cell ; 3 and 5 strongly stalked, 
4 absent ; 6, 7 stalked ; 8 anastomosing with 7. 

(1) *Navasota hebetella, 

Navasota hebetella Rag. N. Am. Phyc. p. 18 (1887); id. Rom. 
Mem. viii. p. 369, pi. 39. f. 5 ; Dyar, Cat. Lep. K. Am. p. 439. 
U.S.A., Texas. 

(2) tNAVASOTA persectella, sp. n. 

$ . Head and thorax purplish red ; abdomen white, dorsally 
sufi'used with fulvoias yellow towards base ; pectus and legs 
white tinged with rufous. Fore wing purplish pink ; the costal 
area creamy white, narrowing to a point at apex, vein 12 defined 
by the costa above it being purplish red and by a fine streak 
below, the costal area defined by a red-bi'own fascia below ; 
veins 4, 3, and 1 slightly streaked with white. Hind wing white 
slightly tinged with ochreous brown. 

Timor, Dili {Boherty), 1 $ type. Exp. 16 mm. 

(3) tNAVASOTA H/EMAPH^ELLA, sp. n. 

5 . Head and thorax dusky crimson ; abdomen fuscous, the 
base of dorsal area fulvous. Fore wing dusky crimson with a 
slightly defined rather paler costal fascia attenuated to a point 
befo]'e apex ; a very slight irroration of black scales and traces of 
a terminal series of points. Hind wing ochreous tinged with 
pale fuscous. 

LouisiADES, St. Aignan I. [Meek), 1 5 type. Exjj. 24 mm. 

(4) tNAVASOTA LEUCONEURELLA, sp. n. 

J . Head and thorax dusky crimson ; abdomen ochreous 
white. Fore wing crimson ; the veins streaked with white, the 
subcostal and median nervures strongly so and the former 
defined on lower side by brown saffusion from base to apex. 
Hind wing yellowish white, the costal area tinged with fuscous. 

Uganda, Kampala (Ansorge) ; Transvaal, White R. (Cooke), 
1 c? ; Natal, Estcourt (Hutchinson), 1 c? type ; Cape Colony, 
Transkei (Miss F. Barrett), 1 S . Exp. 24 mm. 

(5) tNAVASOTA CHIONOPHLEBIA, Sp. n. 

Head and thorax purplish i-ed mixed with some ochreous 
white ; abdomen rufous, dorsally fulvous yellow on basal half ; 
pectus and legs whitish sufiused with rufous. Fore wing pui-plish 
pink, the veins strongly streaked with white. Hind wing white 
tinsred with rufous. 



PYRALID.E, SUBFAMILY HYPSOTROPIN^. 67 

S. Nigeria, Yorubalan<^, Ogbomoso (Sir G. Carter), 1 5 ; 
IST. Nigeria, Zungern (Simpson, Macfie), 2 § , Minna (Macfe), 
1 J, 1 $ type. Exp. 16-22 mm. 

(6) flSTAVASOTA SYRIGGIA, Sp. n. 

$ . Head and thorax rufous tinged with purplish pink, the 
metathorax ochi-eous wliite ; abdomen fulvous yellow ; pectus 
■ and legs white, the latter tinged with purplish rufous. Fore 
wing pale purplish pink, the veins slightly streaked with white. 
Hind wing- white with a faint ochreous tinge. 

Argentusta, Tucuman, Los Vasquez (Dinelly), 1 § type. Exjy. 
24 mm, 

(7) Navasota discipunotella, sp. n. 

5 . Head, thorax, and abdomen white mixed with brown, the 
last faintly tinged with fulvous yellow towards base of dorsum. 
Fore wing white irrorated with fuscous brown and some pale 
purplish-pink scales; a black discoid al point; a slight blackisli 
j)Ostmedial shade from vein 2 to inner margin. 

IST. Nigeria, Minna (Macfie), 1 2 type. Exp. 20 mm. 

A'uctorum. 

Navasota mi/riolecta Dyar, Pr. U.S. Nat. Mus. xlvii. p. 347 
(1913) Panama. 

Genus Hypsotbopa. 

■J'ype- 
Hypsotropa Zell. Isis, 1 848, p. 59 1 limhetla. 

Heospliora Meyr. P. Linn. Soc. N.S.W. vii. 

p. 158 (1882) psamathella. 

Seleucia Jiag. Ann. Soc. Ent. Fr. 1887, p. 259. semirosella. 

Peoria Pi,ag. N. Am. Phyc. p. 19 (1887) approximella. 

Amhala Rag. Nouv. Gen. p. 45 (1888) fascostrigella. 

Lymira Rag. Nouv. Gen. p. 46 (1888) semirosella. 

Tiarra Rag. Nouv. Gen. p. 46 (1888) pusiUella. 

Socora Rag. Nouv. Gen. p. 46 (1888) tenuicostella. 

Talamha Rag. Nouv. Gen. p. 47 (1888) tenuinervella. 

Wekiva Hulst, Trans. Am. Ent. Soc. xvii. 

p. 215 (1890) lateicostella. 

Tinitinoa Dyar, Pr. U.S. Nat. Mus. xlvii. 

p. 347 (1913) phyrdes. 

Proboscis aborted and minute ; palpi typically obliquely up- 
turned to far above vertex of head, the 2nd joint fringed with 
scales in front ; maxillary palpi slightly dilated with scales ; 
frons smooth, with tuft of scales ; antennte of male typically 
somewhat laminate and ciliated, the shaft slightly curved ;it 
base. Fore wing narrow, the apex rounded ; vein 2 from towards 
angle of cell ; 3 and 5 separate, 4 absent ; 6 from below upper 
■angle; 8, 9, 10 stalked, 8 from beyond 10; 11 from cell. Hind 

5* 



68 SIR GEORGE HAMPSON ON THE 

wing with vein 2 from just before angle of cell ; 3 and 5 strongly 
stalked, 4 absent ; 6, 7 shortly stalked ; 8 strongly anastomosing 
with 7. 

Sect. I. Palpi of male uptuvned. 

A. Palpi of male hollowed out to contain the brush-like maxillary palpi; the 
auteunaj with sinus and ridge of scales at base of shaft. 
a. {Amhctla). Antonnaj of male with rather long uniseriate branches, the apex 
ciliated. 

(1) *Hypsotropa contrastella. 

Amhala contrastella Rag. ISTouv. Gen. p. 45 (1888); id. Rom. 
Mem. viii. p. 370, pi. 38. f. 11. 

Natal. 

(2) Hypsotropa fuscostrigella. 

Amhala fuscostrigella Rag. ISTouv. Gen, p. 45 (1888) ; id. Rom.. 
Mem, viii. p. 371, pi. 38. f. 10. 
Punjab. 

(3) Hypsotropa subcostella, sp, n. 

Head and thorax white, the tegulse and patagia suft'used with 
red-brown; antennfe red-brown ; palpi irrorated with red- brown ; 
abdomen white, usually irrorated with brown and dorsally suf- 
fused with fulvous yellow towards base ; legs irrorated with 
brown. Fore Aving white with a faint pink tinge, and strongly 
irrorated with brown ; an ochreous-brown fascia through the 
cell from base to apex, extending to just below the cell. Hind 
wing white with a faint ochreous tinge. 

N. Nigeria, Zungeru (Mac£e, Simpson), 1 J , 2 5 type ; 
Br. E, Africa, K. Kavirondo, Maramas Distr., Ilala [jVeave),. 
2 d . Ex]). 20-28 mm. 

(4) Hypsotropa pervittella, sp. n. 

Head and thorax white mixed with red-brown ; abdomen 
white tinged with red-brown and dorsally sufl'used with fulvous 
yellow towards base. Fore wing white thickly irrorated with 
red-brown ; a red-brown fascia through the cell from base to 
apex, the veins below it streaked with white. Hind wing white 
with the costa tinged with brown towards apex. 

Punjab, Ajmere, 1 c5' ; Bombay, Deesa (jYurse), 1 J , 1 $ 
type. Exp. 20 mm. 

b. Antenna3 of male with uniseriate branches below the sinus onl\', then 
serrate. 

(5) Hypsotropa heterocerella. 

ffi/jysotropa heterocerella Hmpsn. Moths Ind. iv. p. 54 (1896);; 
id. Rom. Mem, viii. p. 371, pi. 52. f. 1. 
Kashmir ; Punjab. 



PYRALID.E, SUBFAMILY HYPSOTROPIN.^. 69 

c. Antemiaa of male somewliat laniinate and ciliated to base. 

(6) HypSOTROPA OCRACEELLA, sp. 11. 

(S . Head and thorax brownish ochreous mixed with some 
white ; abdomen ochreous white, dorsally tinged with fulvous 
yellow towards base. Fore wing brownish ochreous ; a dark 
brown fascia along median nervure ; a diffused anteme'lial line 
from cell to inner margin ; a diffused dark postmedial line tend- 
ing to form streaks in the interspaces, oblique to vein 6, then 
inwardly oblicjue ; a slight dark terminal line. Hind wing 
white tinged \yith ochreous brown and with a dark terminal line 
to submedian fold. 

Timor, Oinaiiiisa (Doherty), 1 c? type. Exj). 14 mm. 

B. Maxillary palpi of male filiform. 

a. Aiiteiuia; of male with sinus at base of sliaft containino- a ridge of soales. 
a' . (Socora). Palpi of male with the 2nd joint obliquelj' upturned and thiclcli' 
scaled, the third joint porrect. 

(7) *Hypsotropa TENUICOSTELLA. 

Socora tenuicostella Rag. Nouv, Gen. p. 46 (1888); id. Rom.- 
Mem. viii. p. 374, pi. 39. f. 6. 
Gambia. 

h' . (Lj/mii-a). Palpi of male slender, the 3rd joint upturned. 

(8) *Hypsotropa SEMIROSELLA. 

Seleucia semirosella Rag. Ann. Soc. Ent. Fr. 1887, p. 259; id. 
Rom. Mem. viil p. 373, pi. 38. f. 9; Stand. Cat. Lep. pal. ii. 
p. 83. 

Syria. 

(9) Hypsotropa illectalis. 

Addyme illectalis Wlk. xxx. 959 (1864); Rag. Rom. Mem. viii. 
p. 373," pi. 38. f . 8. 

Seleucia costatella Rag. Nouy. Gen. p. 46 (1888). 

Borneo ; Celebes. 

(10) tHYPSOTROPA BISCRENSIS, Sp. 11, 

Head and thorax white' tinged with red-brown ; abdomen 
creamy white, dorsally suffused with ochreous brown towards 
base. Fore wing white tinged with ochreous except a white 
costal fascia narrowing to a point at apex, defined below by a 
red-brown fascia ; the median nervure and veins 5, 3 slightly 
streaked with white. Hind wing creamy white. 

Algeria, Biskra {Walsingham), 1 d' , 2 $ type, Hammam-es- 
Salahin (Walsivgham), 1 S • Exp. 22-28 mm. 

(11) tHYPSOTROPA RHODOCHKOELLA, Sp. n. 

Head and thorax deep rose-red ; abdomen ochreous yellow, 



70 SIR GEORGE HAMPSON ON THE 

doraally fulvous yellow towards base ; antennse fulvous yellow ;■ 
pectus, legs, and ventral surface of abdonien ochreous yellow 
suffused witli brown. Fore wing deep i-ose-red, the costa and 
veins strongly streaked with white, the interspaces of costal 
area tinged with bi-own. Hind wing ochi'eous white slightly 
tinged with bi'own. 

Hab. Uganda, Mulema [Doggett), 1 c5' , Ketoma (Doggeii), 

1 c? ; Br. 0. Africa, Mt. Mlanje (A^eave), 7 c? , 3 $ type ; Trans- 
vaal, White R. (Cooke), 1 (S ; Natal, Estcourt {Hictchinson), 

2 c? . I^x]). 20-28 mm. 

(12) tHYPSOTROPA ALBIVENALIS, 

Amhala albivenalis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi. 
p. 1250(1912). 
Ceylon. 

I). Autennaj of male without sinus at base of shaft containing a ridge of scales. 
«!. {TIy]}Sotrnpa). Palpi long; antennae of male with the shaft thickened 
at base. 

(13) *Hypsotropa LUTBICOSTELLA. 

Rypsotropa luteicostella Rag. N. Am. Phyc. p. 19 (1887) ; id. 
Rom. Mem. viii. p. 376, pi. 39. f. 9 ; Dyar, Cat. Lep. N. Am. 
p. 439. 

U.S.A., Florida. 

(14) tHYPSOTROPA OCHRICOSTELLA, Sp. n. 

5 . Head and tegulaj dark red-bi'own ; thorax purplish r6d ;. 
abdomen fulvous yellow ; pectus, legs, and ventral surface of 
abdomen red-brown. Fore wing deep purple-red ; a pale ochreous 
yellow costal fascia narrowing to a point at apex, the costa 
tinged with purple-red and irroi'ated with a few dark scales to 
beyond middle, the costal fascia defined by difl'used dark brown 
below. Hind wing reddish brown. 

Masiionaland (Dobbie), I $ type. JSx]}. 20 mm. 

(15) tHYPSOTROPA CHIONORHABDA, sp. n. 

Head and thorax black-brown mixed with some purple-grey, 
especially on dorsum of thorax ; antennae whitish tinged with red- 
brown ; abdomen pale reddish brown with darker segmental 
lines ; pectus, legs, and ventral surface of abdomen grey suffused 
with blackish, the tibiae and tarsi in front white. Fore wing- 
purplish grey irrorated with black-brown ; a silvery white costal 
fascia narrowing to a point at apex and leaving the costal edge 
black towards base, defined below by a broad black-brown fascia 
with diffused lower edge. Hind wing grey suffused with reddish- 
blown and with darker terminal line. 

S. Nigeria, Yorubaland (Sir G.' Carter), IS, 1 ? ; N. 
Nigeria, Zungeru {Macjie), 1 S ; Uganda, Gondokoro [Reymes- 
Cole), 1 S , Ketoma {Doggett), 2 J ; Br. C. Africa, Mt. Mlanje 



PYRALID.E, SUBFAMILY HYPSOTUOPIN^. 71 

{Keave), 5 cT , 3 $ , Luchenya R. [Xeave), 6 d" , 2 $ type, Ruo 
Valley {Keave), 1 J. Exp. 16-24 mm. 

(16) Hypsotropa infumatella. 

Hypsotropa ivfitmatella Hmpsn. Rom. Mem. viii. p. 377, pi. 39. 
f. 8(1901). 

Transvaal ; ISTatal. 

(17) *Hypsotropa unipunctella. 

Hypsotropa unipunctella Rag. Nouv. Gen. p. 47 (1888) ; id. 
Rom. Mem. viii. p. 377, pi. 38. f. 12 ; Stand. Oat. Lep. pal. ii. p, 12. 
E. Siberia, Amurland. 

(18) f Hypsotropa fusifasciata, sp. n. 

$ . Head and thorax whitish sufiused with red-bl-own ; abdo- 
men fulvous yellow, the ventral sviiface white tinged with red- 
l:)rown. Fore wing whitish tinged with red-brown and irrorated 
with blackish ; a white costal fascia irrorated with blackish,, 
rather diffused below, the costal eage blackish towards base ; 
a black discoidal point ; the terminal area suffused with blackish 
except at costa ; cilia with a blackish line near base. Hind- 
wing whitish strongly suffused with brown ; a fine dark terminal 
line and line near base of cilia. 

Ceylon, Kandy {Green), 1 5 type. Ea'j). 18 mm. 

(19) *Hypsotropa zophopleura. 

Hypsotropha zophopleura/ Turner, Pr. R. Soc. Queensl. xviii.. 
p. 117 (1903). 
Queensland. 

(20) tHYPSOTRUPA PURPURELLA, Sp. n. 

Head and thoi^ax purple-pink ; antennfe fulvous yellow ; palpi^ 
tinged with fuscous ; abdomen fulvous yellow ; pectus, legs, and 
ventral sui'face of abdomen greyish suffused with fuscous, the 
claspers of male white. Fore wing purple-pink irrorated with 
black, the costa, brownish; a black spot in the cell near base; 
antemedial spots in the cell and on vein 1 ; obliquely placed 
spots at upper and loAver angles of cell ; a subterminal series of 
four spots, the spot at discal fold further from termen ; a terminal 
series of black points. Hind wing ochreous white ; a terminal 
black line ; cilia blackish at tips. 

Br. C. Africa, Mt. Mlanje {yeave), 9 d , 3 5 type, Exp., 
d 20, 2 22 mm. ' 

(21)' Hypsotropa limbella. 

Hypsotropa Umhella Zell. Isis, 1848, p. 591 ; Herr.-Schaff. 
Eur. Schmett. iv. p. 110, Tin. f, 38; Rag. Rom. Mem. viii. 
p. 376 ; Stand. Oat. Lep. pal. ii. p. 12. 

S. France ; Italy ; Dalmatia : Asia Minor. 



72 



SIR GEORGE HAMPSON ON THE 



(22) Hypsotropa syriacella, 

Ilypsotropa syriacella Rag. Nouv. Gen. -p. 46 (1888) ; id. Rom. 
Mem. viii. p. 377, pi. 39. f. 7 ; Staud. Cab. Lep. pal. ii. p. 12. 
Syria. 

(23) *Hypsotropa solipu^ctella. 

Hypsotropa solipimctella Rag. Rom, Mem. viii. p. 377, pi. 43. 
f. 23(1901). 
Japan. 

(24) *Hypsotropa paucipunctella. 

Hy2Jsotropa paucipimctella Rag. Bull. Soc. Eat. Fr. 1895, 
p. cii; id. Rom. Mem. viii. p. 378, pi. 51. f. 10 ; Stand. Cat. Lep. 
pal. ii. p. 13. 

Asia Minor, Taurus. 

(25) *Hypsotropa ichorella. 

Anerastia ichorella Led. Vevh. zool.-bot. Ges. Wien, 1855, 
p. 221, pi. 3. f. 8; Ra,g. Rom. Mem. viii. p. 378, pi. 39. f. 10; 
Staud. Cat. Lep. pal. p. 13. 

Syria. 

(26) Hypsotropa vulneratella. 

Epischnia vulneratella Zell. Isis, 1847, p. 769 : Rag. Rom. 
Mem. viii. p. 378, pi. 39. f. 11 ; Staud. Cat. Lep. pal. ii. p. 13. 

Anerastia ostrinella Lah. Bull. Soc. Vaud. vi. Conti\ Faun. Sic. 
p. 396 (1861). 

Hypsotropa roseostrigella Rag. Rom. Mem. viii. p. 379, pi. 39. 
f. 12 (1901). 

Sardinia ; Sicily ; Dalmatia ; Syria. 

(27) fHYPSOTROPA sceletella. 

Anerastia sceletella Zell. Stett. Ent. Zeit. 1867, p. 404 ; Hmjisn. 
Moths Lid. iv, p. 54; Rag. Rom, Mem. viii. p. 379, pi. 44. f. 22. 
Bengal, Calcutta. 

(28) Hypsotropa vertheimsteini. 

Hyp)sotropa ivertheimsteini Rebel, Rovart. Lep. xx. p. 171 
(1913). 

Hungary, 

(29) fhypsotropa punctinervella, sp, u, 

Head and thorax pale red mixed with whitish ; abdomen 
creamy white, dorsally fulvous yellow towards base ; palpi rufous, 
white below towards base : pectus and legs creamy white, the latter 
irroiated with some red-brown. Fore wing pale i-ed slightly 



PYRALID.E, SUBFAMILY HYPSOTROPIN.^?. 73 

irrorated witli dark brown, the veins and costal edge white; a 
minute antemedial black spot on vein 1 ;ind postinedial spots on 
veins 3, 2, 1 ; a terminal series of sliglit blackish points. Hind 
wing ochreous white. 

Ceylon, Haputale (Alston), 1 6 type, Kegalle (Alston), 1 2 , 
Hambantota {Pole), 1 ? . Exj). 14-16 mm. 

(30) Hypsoteopa adumbratella. 

Hypsotropa adumbratella Rag. Nouv. Gen. p. 47 (1888); id. 
Rom. Mem. viii. p. 380, pi. 38. f. 13. 
Gambia ; Natal ; Cape Coloxy. 

h'. (Tiarra). Palpi short; anteniise of male with the shaft curved at base, 

(31) *Hypsotropa pusillella. 

Tiarra piisillella Rag. Nouv. Gen. p. 46 (1888) ; id. Rom. Mem. 
v'iii. p. 372, pi. 39. f . 4''. 
Zanzibar. 

Sect. II. Palpi of male porrect. 

A. (Talamha). Palpi of male hollowed out to receive the brush-like maxillary 
palpi; antenna} with sinus at base of shaft containing a ridge of scales. 

(32) Hypsotropa tenuinervella. 

Talamha tenvAnervella Rag. Nouv. Gen. p. 47 (1888) : id. Rom. 
Mem. viii. p. 387, pi. 40. f. 25 ; Hmpsn. Moths Ind. iv. p. 55. 
Punjab, Kangr^ ; Madras, Nilgiris. 

B. Maxillary palpi filiform. 

a. Antennae of male with sinus at base of shaft containing a ridge of scales. 
fli. {Tinitinoa). Antennae of male pectinate with long uniseriate branches 
to middle. 

(33) *Hypsotropa phyrdes. 

Tinitinoa 'phyrcUs Dyar, Pr. U.S. Nat. Mus. xlvii. p. 348 
(1913). 
Panama. 

h' . (Heosflwra). Antennas of male laminate. 

(34) *Hypsotropa papuasella. 

Reosphora papuasella Rag. Nouv. Gen. p. 47 (1888) ; id. Rom. 
Mem. xxiii. p. 382, pi. 39. f. 14. 
N. Guinea. 

(35) *Hypsotropa sabuletella. 

Anerastia sahuletellaZeW.'LB^. Micr. Caffr. p. 73 (1852); Rag. 
Rom. Mem. viii. p. 383, pi. 39. f. 18. 
Cape Colony. 



74 SIR GEORGE HAMPSON ON THE 

(36) tHypsoTROPA cremoricosta, sp. n. 

J . Head and thorax ochreous, the head and tegulpe suffused 
with purplish red ; antennse ochreous yellow ; abdomen ochreous, 
dorsally fulvous yellow towards base ; pectus, legs, and ventral 
surface of abdomen purplish red. Fore wing pale flesh-red, 
deepening in colour towards the costal fascia which is ochreous 
white, narrowing to a point at apex, slightly defined by brown 
below and leaving the costal edge brown towards base. Hind 
wing ochreous white, the costal area and termen except towards, 
tornus deeper ochreous. 

U.S.A., Colorado, Colorado Springs [Cockerell), 1 (5 type. 
Bxp, 24 mm. 

(37) tHYPSOTROPA NIVEICOSTA, Sp. n. 

$ . Head and thoi-ax white suffused with rufous except on 
dorsum of thorax : abdomen ochreous white, dorsally fulvous 
yellow towards base ; pectus, legs, and ventral surface of abdomen 
white suffused with rufous. Fore wing pale flesh-red irrorated 
with a few red-brown scales and deepening to rufous towards 
the costal fascia, which is snow-white narrowing to a point at 
apex and leaving the costal edge rufous towards base. Hind 
wing white with a faint ochreous tinge. 

Argentina, Gran Chaco, Florenzia (Wagner), 1 $ type. £Jap. 
20 mm. 

(38) tHyPSOTROPA PERIPHiEA, sp. n. 

5 . Head and tegula? white tinged \\ith red-brown ; thor'ax 
pale piuplish pink ; abdomen white dorsally tinged with fulvous 
yellow towards base; pectus, legs, and ventral surface of abdomen 
white tinged with red-brown. Fore wing pale purplish pink ; a 
pure white costal fascia irx'orated with a few brown scales towards 
costa, narrowing to a point at apex and defined below by dark 
red-brown, diffused below. Hind wing white with a faint 
ochreous tinge. 

N. Nigeria, Minna (Macfie), 2 § type, Borgu, Yelwa Lake 
(Migeod), 1 $. Ewp. 16 mm. 

(39) tHYPSOTROPA TRIPARTALIS, Sp. n. 

5 . Head white suffused with rufous ; thoi-ax pale purplish 
pink ; abdomen ochreous yellow ; pectus and legs ochreous 
yellow, the foi'e legs tinged with browii. Fore wing with narrow 
pure white costal fascia leaving the costal edge brown towards 
base, the area to discal fold red- brown and the area below it 
bright purplish pink. Hind wing white with a faint ochreous 
tinge. 

Formosa, Takow {Wileman), 1 $ type. Exp. 18 mm. 

(40) tHYPSOTROPA LATERCULELLA. 

Anerastia laterculella Zell. Stett. Ent. Zeit. 1867, p. 403;. 



PYRAIilD.E, SUBFAMILY HYPSOTROPIN^. 75- 

Hnipsn, Motlis Intl. iv, p. 55 ; Rag. Rom. Mem. viii. p. 383^ 
p].44. f. 21. 
Bengal. 

(41) tHYPSOTROPA ROSESCENS, »Sp. 11. 

(5 . Head and thorax purplish red-br-own, the latter with the 
dorsum grey-brown ; abdomen fulvous yellow ; pectus, legs, and 
ventral surface of abdomen whitish tinged with brown. Fore 
wing pale grey-brown sufiused with crimson-red, rather browner 
towards the costal fascia, Avhich is white slightly irrorated with 
pale I'ed and narrowing to apex. Hind wing white tinged witL 
ochreous brown. 

Ceylox, Xawalapitiya (Green), 1 c? type. Ux]}. 20 mm. 

(42) *Hypsotropa stereosticha. 

Hypsotropha stereosticha Turner, Pr. R. Soc. Queens!, xix. 
p. 4i (1905). 

QuEENSLAXD, Thursday I. 

(43) Hypsotropa icasmopis. 

HypsotropJta icasmojjls Turner, Pr. R. Soc. Queensl. xviii. p. 116. 
(1903). 

queenslais'd. 

(44) Hypsotropa euryzona. 

Hypsotrojxo euryzona Meyr. Ent. Mo. Mag. xix. p. 256 (1882) ; 
Rag. Rom. Mem.viii. p. 382, pi. 39. f. 13. 
S. Australia. 

(45) Hypsotropa xiphopleura. 

Hypsotropha niphopleura Turner, Pr. R. Soc. Queensl. xxiv. 
p. Ill (1913). 

N. Australia. 

(46) tHYPSOTROPA DIAPH.EA, SJ3. n. 

5 . Head and thorax black-brown, the frons and metathorax 
rufous; antennae rufous; palpi black with a greyish gloss; 
abdomen grey suffused with brown; pectus, legs, and ventral 
surface of abdomen grey suffused with blackish. Fore win^ 
whitish ; the costal edge black with a flesli-red fascia below it to 
beyond middle, the area below the cell black Avith a whitish 
streak along vein 1. Hind wing whitish strongly suffused with 
fuscous brown, the cilia whiter with a dark line near base. 

Br. C. Africa, Mt. Mlanje {Neave^j, 1 $ type. Exp. 24 mm. 

(47) Hypsotropa dyseimata. 

Hypsotropha dyseimata Tui-ner, Pi-. R. Soc. Queensl. xxiv.. 
p. 112 (1913). 

Timor Laut ; X. Australia. 



76 sir georae hampson on the 

(48) Hypsotropa quadripunctella. 

H ijpsotropa quadripunctdla Huipsu. J. Bomb. Nat. Hist. Soc. 
xii. p. 307 (1897) ; id. Rom. Mem. viii. p. 384, pi. 39. f. 16. ' 
Punjab, Oude ; Borneo ; Pulo Laut. 

(49) Hypsotropa psamathella. 

Anerastia 2^sainathella Meyi-. Proc. Linn. Soc. N.8.W. iv. 
p. 234 (1879) ; Rag. Rom. Mem. viii. p. 384, pi. 39. f. 15. 
"f Anerastia nitens Butl. Trans. Ent. 8oc. 1886, p. 440. 
Queensland ; N. S. Wales. 

(50) tHYPSOTROPA MONOSTIDZA, Sp. n. 

$ . Head and thorax oclireons with a faint purplish pink 
tinge ; abdomen ochreous with a, fulvous yellow tinge ; pectus, 
legs, and ventral surface of abdomen ochreous with a l:>rownish 
tinge. Fore wing ochreous faintly tinged with purplish pink 
■ and irrorated with dark brown ; a minute blackish discoidal spot. 
Hind wing white tinged with ochreous especially towards termen. 

Sierra Leone {Clements)^ 1 9 type. Exj). 18 mm. 

(51) tHYPSOTROPA GRAPTOPHLEBIA, Sp. n. 

(S . Head and thorax white mixed with purplish red -brown ; 
abdomen white slightly tinged with browu ; pectus, legs, and 
ventral surface of abdomen white suffused with red-brown. 
Fore wing white suffused with pui'plish red-brown and slightly 
irrorated with dark brown, the veins wliite defined on each side 
by fine dark brown streaks. Hind wing white tinged with 
ochreous brown. 

Mashonaland, Salisbury {Marshall), 1 6 type. Exp. 20 mm. 

(52) tHYPSOTROPA POLYACTINIA. 

Heosjihora polyactinia Hmpsn. Rom. Mem. viii. p. 384, pi, 52. 
f. 2 (1901). 

Uganda ; Mashonaland ; Transvaal ; ISTatal. 

(53) tHYPSOTROPA ENDORHODA, Sp. U. 

$ . Head and thorax ochreous suffused with purplish pink ; 
abdomen ochreous white, dorsally fulvous yellow towards base ; 
pectus, legs, and ventral surface of abdomen ochreous white 
suffused with red-brown. Fore Aving with the costal half to 
median nervure and vein 5 ochreous, tinged with purplish pink 
towards costa and with slight pale streaks on the veins ; the 
inner half purplish pink with slight white streaks on the veins. 
Hind wing white tinged with ochreous. 

N. Rhodesia {Coryndon), 1 2 type. Exp. 24 mm. 

(54) Hypsotropa ramulosella. 
Heosphoraramulosella^ii.g.^u\\. Soc. Ent. Fr. 1895, p. cii; id. 



PYRALIDiE, SUBFAMILY HYPSOTROPINyE. 77 

Rom. Mem. viii. p. 385, pi. 52. f. 3 ; Stand. Cat. Lep. pal. ii. 
p. 12. 
Sykia. 

(55) Hypsotkopa leucophlebiella. 

Heosphora leucophlebiella Rag. Nouv. Gen. p. 47 (1888); id. 
Rom. Mem. viii. p. 385, pi. 39.1 17. 

Br. C. Africa ; Mashonaland ; Transvaal ; Natal ; Cape 
Colony. 

(56) Hypsotropa rhodosticha. 

Hypsotropha rhodosticha Tnrner, Pr. R. Soc. Qneensl. xviii. . 
p. 116 (1903). 
Queensland. 

h. {Peoria). AntenuEe of male without sinus and ridge of scales at base of shaft 
•(57) tHYPSOTROPA POLYSTICTELLA, sp. D. 

c7 . Head, thorax, and abdomen whitish suffused with rufous. 
Fore wing ochreous white suffused with rufous in the inter- 
spaces leaving the veins pale ; the costal edge black towards 
base ; a slight shade formed by black irroration below the costa 
to apex ; a small antemedial black spot on vein 1, a small spot 
on lower discocellulars and an oblique postmedial series of four 
small spots on veins 5, 3, 2, 1 ; a terminal series of small black 
spots. Hind Aving white tinged with ochreous ; a fine fuscous 
terminal line and slight brown line near base of cilia. 

Br. C. AFRiCA,J\it. Mlanje (JS^eave), 2 d type. Uxp. 20 mm. 

(58) *Hypsotropa acnidias. 

Hypsotropha acnidias Turner, Pr. R. Soc. Queens!, xviii. p. 117 
(1903). 

Queensland. 

(59) tHYPSOTROPA approximella. 

Eurhodo'pe appj-oximella Wlk. xxxv. 1722 (1866); Rag. Rom. 
Mem. viii. p. 386. pi. 40. f. 24 ; Dyar, Cat. Lep. N. Am. p. 439. 

Anerastia hcematica Zell. Yerh. zool.-bot. Ges. Wien, 1872, . 
p". 555. 

Anerastia roseatella Pack. Ann. N. Y. Lye. ISTat. Hist. x. 
p. 270 (1870). 

U.S.A., Massachusetts, IS". York, Pennsylvania, Ohio, Illinois, 
N. Carolina, Texas, Colorado. 

(60) tHYPSOTROPA LEUCOCRASPIS, Sp. n. 

d . Head, thorax, and abdomen white suffused with ochreous 
brown. Fore wing pale ochreous brown, the costal edge white. 
Hind wing white tinged with ochreous bi'own especially on apical 
area. 

Argentina, Corrientes, Goya (Perrens), 1 cJ type. Ex2}. 20 mm. . 



78 sir george hampson on the 

(61) *Hypsotropa bipartella, 

Peoria hipartella Rag. N". Am. Phyc. p. 19 (1887) ; id. Rom. 
Mem. viii. p. 386, pi. 4U. f. 33 ; Dyar, Cat. Lep. N. Am. p. 439. 
U.S.A., N. Carolina. 

Auctorum. 

Peoria albidella Hnlst, Can. Ent. xxxiii. p. 175 (1900); Dyar, 

Cat. Lep. K Am. p. 439 U.S.A., California. 

llyjjsotropha larojyis Turner, Pr. R. Soc. Queensl. xxiv. p. 173 

(1913) Queensland. 

,, neurica Turner, Pr. R. Soc. Queensl. xxiv. p. 113 

(1913) N.Australia.. 

,, castella Pag. Zoologica., xxix. ^. 163 (1900). 

Bismarck Arch. 

Genus Raphimetopus, nov. 

Type, R. spinifrontella. 

Proboscis aborted and minute ; palpi downcurved, extending 
about three times length of head and thickly scaled ; maxillary 
palpi minute and filiform ; frons with long flattened corneous 
plate, somewhat trifid in front and produced to short lateral 
processes, a corneous plate below the frons; antennpe of male 
laminate. Fore wing narrow, the apex rounded, the termen 
obliquely curved ; vein 2 from close to angle of cell ; 4 absent ; 
5 from above angle ; 6 from below upper angle ; 8, 9 stalked ; 
10, 11 from cell. Hind wing with vein 2 from just before angle 
of cell ; 3 and 5 stalked, 4 absent; 6, 7 shortly stalked ; 8 closely 
approximated to or anastomosing with 7. 

(1) Raphimetopus spinifrontella. 

Anerastia spinifrontella Rag. Nouv, Gen. p. 48 (1888) ; id. 
Rom. Mem. viii. p. 399, pi. 40. f. 2 ; Hmpsn. Moths Ind. ii. 
p. 56. 

Bombay, Sind. 

(2) Raphimetopus ablutella. 

Anerastia ablutella Zell. Isis, 1839, p. 17; Herr.-Schaff. Eur. 
Schmett. iv. p. 109, Tin. f. 89; Rag. Rom. Mem. viii. p. 403; 
Hmpsn. Moths Ind. iv. p. 56 ; Staud. Cat. Lep. pal. ii. p. 12. 

f Anerastia stigmatella Rag. Nouv. Gen. "p. 49 (1888) ; id. Rom. 
Mem. viii. p. 403, pi. 40. f. 4. 

Anerastia himaculata Rag. Nouv. Gen. p. 49 (1888) ; id. Rom. 
Mem. viii.' p. 404, pi. 40. f. 5 ; Hmpsn. Moths Ind. iv. p. 56. 

Anerastia niajorella Roths. Nov. Zool. xx. p. 138 (1913). 

Spain ; Sicily ; Algeria ; N. Nigeria ; Congo, Egypt ; Da- 
maraland; Transvaal; Orange R. Colony; Aden; Asia Minor; 
Persia; W.Turkestan; Japan; Kashmir; Punjab; Madras. 



PYRALID^, SUBFAMILY HYPSOTROPIN.*:. 79 

Auctonmn. 

Anerastia korbi u^adja, Iris, xxiv. p. 117(1910). 

? = /?. abluiella Caspian Sea. 

Genus Metacrateria, nov. 

Type, JI. pidverulella. 

Proboscis absent ; palpi dovvncurved, extending about twice 
the length of head and thickly scaled ; maxillary jDalpi minute 
and filiform ; frons with long truncate corneous prominence with 
raised rim at extremity, a flattened plate below the frons ; 
antennas of male laminate and ciliated. Fore wing rather 
narrow, the apex rounded, the termen evenly curved ; vein 2 from 
near angle of cell ; 4 absent ; 5 from above angle ; 6 from below 
upper angle; 8, 9 stalked; 10, 11 from cell. Hind wing with 
vein 2 from just before angle of cell ; 3 and 5 stalked, 4 absent ; 
6,. 7 shortly stalked ; 8 approximated to but not anastomosing 
with 7. 

(1) Metacrateria metallactis. 

Anerastia metallactis Meyr. Trans. Ent. Soc. 1887, p. 622 ; 
Rag. Rom. Mem. viii. p. 399, pi. 40. f. 11. 
N. S. Wales. 

(2) Metacrateria pulverulella. s. 

Anerastia pidverulella Hmpsn. Moths Ind. iv. p. 56 (1896) ; 
id. Rom. Mem. viii. p. 400, pi. 52. f. 4. 
Ceylon. 

(3) tMsTACRATERIA PERIRRORELLA, sp. n. 

5 . Head and thorax red-brown tinged with grey and slightly 
irrorated with f uscovis ; abdomen grey suffused with red-brown ; 
palpi below, pectus, legs, and ventral surface of abdomen whitish 
tinged with red-brown. Fore wing whitish suffused with red- 
brown and tliickly irrorated with black ; small spots formed by 
black ii-roration at upper and lower angle of cell ; cilia with a 
whitish line at base. Hind wing whitish strongly suffused with 
reddish brown, the cilia whiter. 

Mashonaland (Bobbie), 2 2 type. Uxj). 28 mm. 

(4) Metacrateria miasticta, sp. n. 

$ . Head, thorax, and abdomen whitish suffused with red- 
brown. Fore wing whitish suffused with red-brown and slightly 
irrorated with black, especially on the veins ; a small black spot 
at lower angle of cell. Hind wing white tinged with red- 
brown. 

Mexico, Presidio {Forrer), 1 5 type, Godman-Salvin Coll. 
Exp. 24 mm. 



80 SIR GEORGE UAMPSON ON THE 

Genus Prinanerastia*. 

Type, /''. lotella. 

Proboscis absent ; palpi down curved, extending about twice 
the len^gth of head and thickly scaled ; niaxilliuy palpi minute 
and filiforru ; frons with rounded prominence with slight raised 
rim at extremity, a corneous plate below the frons ; antennaj of 
male laminate and minutely ciliated. Fore wing narrow, the apex 
rounded, the termen evenly curved ; vein 2 from towards angle of 
cell ; 4 absent ; 5 from above angle ; 6 from below upper angle ; 
8, 9 stalked; 10, 11 from cell. Hind wing with vein 2 from 
just befoi'e angle of cell ; 3 and 5 stalked, 4 absent ; 6; 7 shortly 
stalked ; 8 closely approximated to but not anastomosing with 7.. 

(1) Prinanerastia lactealis. 

Lymire lactealis Koths. Nov. Zool. xx. p. 138 (1913). 
Enosima alhicostalis Roths. Nov., Zool. xx. p. 138 (1913). 
Lymire strictvpennis Roths. Nov. Zool. xxii. p. 237 (1915). 
Algeria ; Tunis. 

(2) Prinanerastia nitidicostella. 

Anerastia nitidicostella Rag. Ann. Soc. Ent. Fr. 1887, p. 259; 
id. Rom. Mem. viii. p. 399, pi. 40. f. 12 ; Stand. Cat. Lep. pal. ii. 
p. 12. 

S. Russia. j 

(3) Prinanerastia lotella. 

Tinea lotella Hiibn. Eur. Schmett., Tin. f. 344 (1817); Dup.. 
Lep. Fr. x. pi. 283. f. 6; Herr.-Schaff. Eur. Schmett. iv. p. 100, 
Tin. m 90-92; Leech, Pyr. pi. 10. f. 3; Rag. Rom. Mem. viii.^ 
p. 397 ; Stand. Cat. Lep. pal. ii. p. 12. 

Tinea miniosella Zinck. Germ. Mag. iii. p. 126 (1818). 

Tinea puherella Hiibn. Eur. Schmett., Tin. f. 454 (1823). 

Britain; France : Germany ; Austria; Hungary; Switzer- 
land ; W. Russia ; Asia Minor ; Persia ; W. Turkestan. 

(4) *Prinanerastia incarnatella. 

Anerastia incarnatella Rag. Ann. Soc. Ent. Fr. 1887, p. 259; 
id. Rom. Mem. viii. p. 398, pi. 38. f. 15 ; Stand. Cat. Lep. pal. ii.. 
p. 12. 

S. Russia. 

Genus Chortonceca, nov. 

Type, C. leucocraspia. 

Proboscis aborted and minute ; palpi of male typically obliquely 
upturned to above vertex of head and thickly scaled, hollowed 
out to receive the brush-like maxillary palpi, of female down- 

* The type of Anerastia Hiibn. is dignella Hiibn. 



PYRALID.E, SUBFAMILY HYrSOTROPIN^. 81 

curved and about twice the length of head ; frons with pointed 
conical prominence; antennae of male typically with rather long 
uniseriate branches, the shaft with sinus at base containing a 
ridge of scales. Fore wing iiariow, the apex rounded, the termen 
evenly curved ; vein 2 from towards angle of cell ; 3 and 5 from 
angle, 4 absent ; 6 from below upper angle ; 8, 9 stalked : 10, 11 
fi'om cell. Hind wing with vein 2 fl-om just before angle of cell ; 
3 and 5 stalked, 4 absent ; 6, 7 shortly stalked ; 8 closely 
approximated to 7 but not anastomosing with it. 

Sect. I. Palpi of male obliquely upturned and hollowed out to receive tlie brush - 
like maxillary palpi ; autennse pectinate with uniseriate branches, the shaft 
with sinus and ridge of scales at base. 

(1) tOnORTONCECA LEUCOCRASPIA, Sp. n. 

Head and thorax whitish suftused with rufous ; abdomen white 
tinged with rufous and dorsally suffused with fulvous yellow 
towards base; palpi white in front; pectus, legs, and ventral 
surface of abdomen white tinged with rufous. Fore wing- 
creamy white slightly tinged with rufous ; a white costal fascia, 
narrowing to a point at apex, slightly irrorated with rufous and 
defined below by a luther diffused rufous streak ; the veins 
white defined on each side by fine rufous streaks ; a fine rufous 
streak in snbmedian fold. Hind wing semihyaline white. 

Algeria, Hammam-es-Salahin {Wcdsingham). 2 cT , 3 5 type. 
Exp. 30-34 mm. 

Larva. White with numei^ous red-brown lines interrupted at 
the incisures, the head and 1st thoracic somite wholly i-ed- 
brown. Feeds in the stems of a reed-like grass. 

Sect. II. Palpi of male obliquelj' upturned and not hollowed out to receive the 
maxillary palpi, which are filiform ; antennse of male laminate and ciliated 
with slight sinus at base of shaft, but no ridge of scales. 

(2) tCHORTONCECA MINIMELLA. 

Meliarpha mmimella Hmpsn. Rom. Mem. viii. p. 392, pi. 52. 
f. 11 (1801). 

Borneo, Pulo Laut ; Celebes, Talaut I. ; N. Australia. 

Sect. III. Palpi of male downcurved and extendirig about three times length of 
head, not hollowed out to receive the maxillary palpi which are filiform ; 
autennaj of male laminate and ciliated, with a sinus at base of shaft con- 
taining a ridge of scales. 

(3) *Chorton(eca minoralis. 

Anerastia minoralis Lower, Tr. R. Soc. S. Austr. 1903, p. 52. 
Queensland. 

(4) Chortonceca eurysticha. 

Anerastia eurysticha Turner, Pr. R. Soc. Queensl. xviii. p. 119 
(1903). 

Proc. Zool. Soc— 1918, No. YI. 6 



82 SIR GEORGE HAMPSON ON TUE 

Hypsotropha niphosema Turner, Pr. R. Soc. Queensl. xxiv. 
p. 112(1913). 
Queensland ; N. Australia. 

Genus Rhinaphe. 

Type. 
Rhinaphe Berg, Bull. Soc. Imp. Nat. Mosc-. xlix. 

2, p. 231 (1874) = signaticollis. 

Ampycophora Meyr. Pr. Linn. Soc. N.S.W. vii. 

p. 158 (1882) cqjotomella. 

Comorta Rag. ISTouv. Gen. p. 48 (1888) itigricostcilis. 

iLTaZiarjj/ia Rag. Nouv. Gen. p. 48 (1888) sep)aratella. 

Homosassa Hulst, Trans. Am. Ent. Soc. xvii. 

p. 214(1890) ella. 

Enosima Rag. Rom. Mem.viii. p. 389 (1901)... neesimella. 
Ampycodes Hmpsn. Rom. Mem. viii. p. 393 

(1901) .* pcdlidicosta. 

Eryth'phlehia Hmpsn. Rom. Mem. viii. p. 393 

(1901) enervella. 

Proboscis absent ; palpi typically dovvncurved, extending about 
tbree times length of head and thickly scaled ; maxillary palpi 
filiform ; frons smooth and rounded ; antennee of male typically 
laminate and ciliated. Fore wing narrow, the apex rounded, the 
termen obliquely curved ; vein 2 from near angle of cell ; 4 
absent ; 5 from above angle ; 6 from below upper angle ; 8, 9 
stalked; 10, 11 from cell. Hind wing with vein 2 from just 
before angle of cell ; 3 and 5 stalked, 4 absent ; 6, 7 shortly 
stalked ; 8 closely approximated to or anastomosing with 7. 

Sect. I. Palpi of male hollowed out to receive the brush-like maxillary palpi. 
A. Palpi of male upturned. 

a. {Ampycophora). Antennse of male uiiiserrate. 

(1) t Rhinaphe approximella, sp. n. 

$ . Head and thorax purplish red, the vertex of head and 
dorsum of thorax whitish tinged with red ; abdomen white, 
dorsally tinged with fulvous yellow towards base ; pectus white ; 
legs red-brown. Fore wing whitish suffused with purplish pink, 
leaving the veins whiter and defined on each side by fine 
purplish-red streaks ; a white costal fascia tinged with purplish 
pink at costa, narrowing to a point at apex, defined below by 
a narrow dark brown streak giving off a fine streak above vein 6. 
Hind wing white, the costal area and termen tinged with brown. 

Queensland, Peak Downs, 1 $ type. Exp. 24 mm. 

(2) Rhinaphe apotomella. 

Pempelia apotomella Meyr. Pr. Linn. Soc. N.S.W. iv. p. 224 
(1879) : Rag. Rom. Mem. viii. p. 388, pi. 40. f. 17. 

Punjab; Ceylon; Selangor; Philippines; Celebes, Sangirl. 
Timor, Oinainisa, Dili ; Queensland. 



PYRALIDyE, SUBFAMILY HYPSOTROPIN.E. 83 

b. (Comorta). Antemise of male laminate. 

(3) *RHI]SrAPHE PLINTHINA. 

Anerastia plinthinci Turner, Pr. R. Soc. Queensl. xix. p. 43 
(1905). 

Queensland ; IST. Australia. 

(4) fRniNAPHE castanealis. 

Anerastia castanealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi. 
p. 1251 (1912). 

Br. C Africa ; Transvaal ; Ceylon ; Bali. 

(5) tBniNAPHE celsella. 

Araxes celsella Wlk. xxvii. 193 (1863) ; Hmpsn. Motlis Ind, 
iv. p. 55 ; Rag. Rom. Mem. viii. p. 405, pi. 40. f . 7. 
Ceylon ; Philippines. 

(6) tRniNAPHE nigricostalis. 

Trachonitis nigricostalis Wlk. xxvii. 40 (1863) ; Hmpsn. 

Moths Ind. iv. p. 57 ; Rag. Rom. Mem. viii. p. 389, pi. 39. f. 22. 
Comorta atricostella Rag. Nouv. Gen. p. 48 (1888)o 
Gambia; S. Nigeria; Transvaal; Ceylon; Burma; Anda- 

MANS; Borneo; Fiji. 

(7) Rhinaphb holoph^a. 

Ampycophora holophma Turner, Pr. R. Soc. Queensl. xix. 
p. 42 (1905). 

Queensland; N. Australia. 

B. Palpi obliquely upturned. 

a. Antennae of male with short uniseriate branches, the apical part ciliated. 

(8) Rhinaphe VECTIFERELLA. 

Enosima vectiferella Rag. Rom. Mem. viii. p. 391, pi. 42. f. 24 
(1901). 

Br. E. Africa ; Uganda ; Br. C. Africa ; Transvaal ; 
Madagascar ; Comoro Is. 

h. (EnosiiJia). Antennse of male laminate. 

(9) * Rhinaphe venella. 

Enosima venella Hmpsn. Rom. Mem. viii. p. 391. pi. 40. f. 19 
(1901). 

Cochin China. 

(10) Rhinaphe neesimella. 

Enosima neesimella Rag. Rom. Mem. viii. p. 390, pi. 43. f. 22 
(1901). 

Japan ; Corea. 

6* 



»^ SIR GEORGE HAMPSON ON THE 

(11) RaiNAPHE FLAVESCENTELLA. 

Enosima flavescentella Hmpsn. Rom. Mem. viii. p. 390, pi. 40. 
f. 18(1901). 

Formosa ; C. China. 

c. (Ampi/codes). Palpi downcurved in both sexes ; antennae vinlserrate and 
ciliated, with a large siuus at base of shaft containing a ridge of scales. 

(12) tRHINAPHE STICTELLA. 

Anerastia stictella Hmpsn. J. Bomb. Nat. Hist. Soc. xviii. 
p. 259 (1908). 
Punjab. 

(13) tRniNAPHE PALLIDICOSTA. 

Anerastia pallididicosta limpsn. Moths Ind. iv. p. 57 (1896) ; 
id. Rom. Mem. viii. p. 393, pi. 39. f. 23. 
Assam ; Burma. 

(14) *Rhinaphe haploschema. 

Ampyco'pTiora h%ploschema Turner, Pr. R. Soc. Queensl. xviii. 
p. 117 (1903). 
Queensland. 



Sect. II. Palpi of male not hollowed out to receive the maxillary palpi, which are 
filiform. 

A. (jilaUarplm) . Palpi of male obliquely upturned ; the antennae laminate and 
ciliated, with a large sinus at base of shaft containing a ridge of scales; 

(15) *Rhinaphe separatella. 

Malimyha separatella Rac^. ISTouv. Gen, p. 48 (1888) ; id. Rom. 
Mem. viii. p. 392, pi. 39. £."21. 
Cameroons. 



B. Palpi downcurved in both sexes. 

a. (lErythphlehia). Antennae of male laminate and ciliated, with a large sinus at 
base of shaft containing a I'idge of scales. 

(16) Rhinaphe enervella. 

' Erythphlehia enervella Hmpsn. Rom, Mem. viii. p. 394, pi. 39. 
f. 24 (1901). 

. N. Guinea ; Louisiade Is. ; Queensland ; W. Australia. 

(17) Rhinaphe virginella. 

Anerastia virginella Meyr. Proc. Linn. Soc. N.S.W. iv. p. 233 
(1879) ; Rag. Rom. Mem, viii. p. 394, pi. 40. f. 8. 
Queensland. 



PYUALID^, SUBFAMILY HYPSOTROPINJ;:. 85 

6. Anteii'ifeof male bipectiiiate, without sinus and ridge of scales at tase of 
shaft. 

(18) RhiNAPHE BISERIELLA. 

Anerastia biseriella Hmpsn. Rom, Mem. viii. p. 397, pi. 52. 
f. 18 (1901). 
Queensland. 

c. [Bhhiapha). Antennje of male laminate and ciliated, without sinus and 
ridge of scales at base of shaft. 

(19) tRniNAPHE HEMIRHODELLA. 

Anerastia hemirhodella Hmpsn. Rom. Mem. viii. p. 402, pi. 52. 
f. 12 (1901). 
S. Brazil. 

(20) Rhinaphe lotricella. 

Anerastia lotricella Zell. Isis, 1848, p. 861 ; Rag. Rom. Mem. 
viii. p. 401, pi. 8. f. 21. 
S. Brazil. 

(21) * Rhinaphe leucot^niella. 

Anerastia kucotceniella Rag. Nouv. Gen. p. 48 (1888) ; id, Rom. 
Mem. viii. p. 401, pi. 40. f. 3. 
Japan. 

(22) tPtHINAPHESANGIRENSIS, Sp. n. 

$ . Head and thorax dark red-brown ; abdomen grey tinged 
with brown and dorsally fulvous yellow towards base ; pectus 
and legs grey suffused with brown. Fore wing deep purple- 
pink ; a white costal fascia narrowing to apex, defined below 
by diffused dark red-brown and with the costal edge and vein 
12 purple-pink. Hind wing whitish tinged with red-brown 
especially on costal area; a fine brown terminal line except 
towards tornus. 

Celebes, Sangir I. (Doherty), 1 $ type. Exp. 18 mm. 

(23) Rhinaphe brunneovittella. 

Anerastia brunneovittella Rag. ISTouv. Gen. p. 49 (1888); id. 
Rom. Mem. viii. p, 401, pi, 40. f. 6 ; Hmpsn. Moths Ind. iv. 
p. 56. 

0. China ; Punjab ; Bombay ; Ceylon. 

(24) tRniNAPHE PH^OSTROTELLA, Sp. n. 

9 . Head and thorax creamy white mixed with red-brown ; 
abdomen creamy white, dorsally fulvous yellow towards base; 
pectus and legs creamy white. Fore wing creamy white irrorated 
with pinkish brown ; a creamy white costal fascia with hardly 



86 SIR GEORGE HAMPSON ON THE 

any brown irroration on it, narrowing to apex and defined below 
by diftused bi'own. Hind wing white. 

Ceylon, Puttalam {Pole), 1 $ type. Exp. 22 mm. 

(25) Rhinaphe syssema. 

Anerastia syssema Turner, Pr. R. Soc. Qaeensl. xxiv. p. 114 
(1913). 

Queensland; IST. Australia ; W.Australia. 

(26) tRniNAPHE lateritiella, sp. n. 

5 . Head and thorax grey-brown mixed with some white ; 
abdomen whitish suffused with red- brown, the basal segment 
white. Fore wing reddish brown, darker towai'ds the pure 
white costal fascia naiTOwing to apex. Hind wing white slightly 
tinged with brown. 

B. Nigeria, Yorubaland, Ogbomoso [Sir G. Carter), 1 $ type. 
ExjJ. 20 mm. 

(27) *RhINAPHE ELLA. 

Ephestia ella Hulst, Ent. Am. iii. p. 138 (1887); Eag. Rom. 
Mem. viii. p. 400, pi. 40. f. 1 ; Dyar, Cat. Lep. N. Am. p. 440. 
U.S.A., Florida. 

(28) Rhinaphe taliella. 

Anerastia taliella Hmpsn. Rom. Mem. viii. p. 402, pi. 53. f. 17 
(1901). 

Queensland. 

(29) fRniNAPHE venilineella, sp. n. 

$ . Head and thorax white irrorated with reddish brown ; 
abdomen white tinged with red-brown, dorsally fulvous .yellow 
towards base. Fore wing white thickly irrorated with reddish 
brown, the veins white defined on each side by fine brown 
streaks. Hind wing white. 

Sudan, Fort Sudan {Waterfield), 1 $ type. Exp. 24 mm. 

(^30) *Rhinaphe infumella. 

Anerastia infumella Rag. Ann. Soc. Ent. Fr. 1887, p. 260; id. 
Rom. Mem. viii. p. 402, pi. 38. f. 16; Staiid. Cat. Lep. pal. ii. 
p. 12. 

Persia. 

(31) *Rhinaphe conspersella. 

Anerastia conspersella Rag. Rom. Mem. viii. p. 404, pi. 40. 
f. 13 (1901). 

U.S.A., Colorado. 



PYRALIDiE, SUBFAMILY UYPSOTROPIS^ 87 

(32) *E.MINAPHE SEBBOLDI. 

Anerastia seeholdi Rag, Aiin. Soc. Ent. Fi". 1894, p. 177 ; id. 
Iris, xi. pi. 1. f. 7; Stand. Cat. Lep. pal. p. 12. 
Spain. 

(33) fRniNAPHE ENDONBPHELE, Sp. n. 

d" . Head, thorax, and abdomen white slightly tinged with 
red-bi'own and irrorated with dark scales, the last dorsally 
fulvous yellow towards base ; antennae brownish. Fore wing- 
white irrorated with blackish and faintly tinged with red-brown ; 
a rather diffused rounded blackish antemedial spot on vein 1 ; a 
terminal series of blackish points. Hind wing white faintly 
tinged with ochreous brown ; a slight brown terminal line. 
Brazil, Rio Janeiro, 2 6 type. Exp. 20 mm. 

(34) tRniNAPHE IGNETINCTA, Sp. n. 

5 . Head and thorax ochreous white suffused with fiery red ; 
abdomen ochreous. Fore wing ochreous suffused with fiery red 
and on costal half tinged with puiplish pink ; oblique diffused 
antemedial and medial blackish bars on inner area ; two slight 
dark discoidal points ; an indistinct oblique brown postmedial 
line ; a terminal series of slight dark points. Hind wing semi- 
hyaline white tinged with ochreous especially towards termen. 

ARGE^fTIls^A, Sta. Fe, Ocampo {Wagner), 2 5 type, Buenos 
Ayres {Wilkinson), 1 5 . Exp. 22 mm, 

(35) *Rhinaphe mictochroella. 

Anerastia mictochroella Rag, ISTouv. Gen, p. 49 (1888) ; id. Rom. 
•Mem. viii. p. 404, pi. 40, f. 10. 
Argentina, Goya. 

(36) tRniNAPHB FURVIMACULA, sp. n. 

$ . Head and thorax ochreous faintly tinged with rufous ; 
abdomen ochreous, dorsally fulvous yellow towards base. Fore 
wing ochreous faintly tinged with rufous and irrorated with 
dark brown scales ; an oVjlique slightly sinuous red-brown almost 
medial line with a conical fulvous-red spot on its outer side 
in submedian interspace ; a small dark brown spot at lower angle 
of cell; an indistinct brown postmedial line, oblique below 
vein 5. Hind wing ochreous white suffused with purple-brown 
especially on terminal half except towards tornus ; cilia whiter. 

Argentina, Tucuman, Los Vasquez {Dinelly), 1 $ type. Ex23. 
28 mm. 

(37) Rhinaphe signicollis. 

Rhina2)he signicollis Berg, Bull. Soc, Imp. Nat. Mosc. xlix. 2, 
p. 23a (1874) ; Rag. Rom. Mem. viii. p. 405, pi. 40. f. 9. 
Argentina, Gran Chaco, Florenzia, Buenos Ayres. 



OO SIR GEOROE IIAMPSON ON THE 

Auctoi'um. 

Anerastia xiphlmela Lower, Tr. R. Soc. S. Austr. 1903, p. 52, 

nr. 7i^, castanealis Queensland. 

„ ahhpta Tuvner, Pr. R. 8oc. Qiieensl. xxiv. p. 114 

(U)lo). i Metacrateria. N.Queensland; 

N. Australia. 
„ argosticha Turner, Pr. R. Soc. Queensl. xxiv. p. 115 

(1913), nr. B. enervella N. Australia. 

,, ancemopsls Turner, Pr. R. Soc. Queensl. xxiv. p. 116- 

(1913), ? nr. R. celsella ' IST. Australia. 

,, biillora Turner, Pr. R. Soc. Queensl. xxiv. p. 116 

(1913), ? nr. R. plinthina Queensland. 

,, acropha'a Turner, Pr. R. Soc. Queensl. xxiv. p. 117 

(1913). '^ Sect. En/thjMebia N, Australia. 

,, pleitrochorda, Turner, Pr. R. Soc. Queensl. xxiv. 

p. 117 (1913). ? Sect. Enosima ... Queensland. 
,, erasmia Turner, Pr. R. Soc. Queensl. xxiv. p. 117 

(1913). ? Sect. Erythpldebia Queensland. 

„ ephesUella Viard, Bull. Soc. Ent. Fr. 1913, p. 82. 

Basses Alpes. 

Genus Sudania. 

Type. 
Siulania Hmpsn. Rom. Mem. viii. p. 380 (1901)... svhcostella. 

Proboscis aborted and minute ; palpi upturned to just above 
vertex of head, the 2nd joint moderatel}' scaled, the 3i'd moderate ; 
maxillary palpi filiform ; frons smooth, with conical tuft of scales ; 
antenna? of male with short uniseriate branches. Fore wing 
rather narrow, the apex rounded ; vein 2 from near angle of cell ; 
3 and 5 from angle, 4 absent; 6 from below npper angle; 8, 9, 
10 stalked, 8 from be^yond 10; 11 from cell. Hind wing with 
vein 2 from before angle of cell ; 3 and 5 from angle, 4 absent ; 
6, 7 stalked ; 8 not anastomosing with 7. 

*Sudania subcostella. 

Sudania subcostella Hmpsn. Rom. Mem. viii. p. 381, pi. 51, 
f. 11(1901). 

CtAEOON. 

Genus Rhodochrysa. 

Type. 
Rhodochrysa Hmpsn. Rom. Mem. viii. p. 387 (1901). superbeUa. 

Proboscis aborted and minute; palpi doAvncurved, extending 
about twice the length of head and thickly scaled ; maxillary 
palpi filiform; frons smooth, with conical tuft of scales; antennfe 
of male laminate and minntely ciliated, the shaft slightly cnrved 
at base but without ridge of scales. Fore wing narrow, the apex 
rounded, the termen obliquely curved ; vein 2 from towards angle 
of cell ; 3 and 5 separate, 4 absent ; 6 from below upper angle ; 
8,9, 10 stalked, 8 from beyond 10; 11 from cell. 'Hind wing 



PYRALID/E, SUBFAMILY UYP.SOTKOPIN.E, 89 

with vein 2 frovn just before angle of cell ; 3 and 5 from angle, 
closely approximated towards origin, 4 absent ; 6, 7 shortly- 
stalked ; 8 closely approximated to 7 but not anastomosing 
with it. 

RHODOCnRYSA SUPEnBELLA. 

Rhodochrysa superbella Hnipsn. Rom. Mem. viii. p. 387, pi. 39. 
f. 12 (1901). 

TuAxsvAAL ; Natal. 

Genns Ccenotropa, nov. 

Type, C. limitella. 

Proboscis absent; palpi downcurved, extending about three 
times length of head and thickly scaled ; maxillary palpi minute 
and filiform ; frons with rounded pi-ominence ; antennae of female 
minutely ciliated. Fore wing long and narrow, the apex rounded ; 
Vein 2 from towards angle of cell ; 4 absent ; 5 from above angle ; 
6 from below upper angle; 8, 10, 11 stalked, 9 absent. Hind 
wing with vein 2 from towards angle of cell ; 3 and 4 absent ; 
6, 7 from upper angle ; 8 anastomosing strongly with 7. 

t Ccenotropa limitella, sp. n. 

2 . Head and thorax red-brown mixed with some flesh-white; 
abdomen red-brown, doisally fulvous towards base. Fore wing 
pale purplish pink; a white costal fascia slightly irrorated with 
purplish red, narrowing to apex and defined below by a rather 
difl'used dark brown streak. Hind wing semihj'aline white tinged 
with flesh-colour, the costal area suffused witli brown. 

Paraguay, Sapucay {Foster), 1 $ type. Exjo. 22 mm. 

Genus Bandera. 

Type. 

Bandera Rag. N. Am. Phyc. p. 19 (188 7) binotella. 

Osceola Hulst, Smith, List N. Am. Lep. p. 85 

(1891), non descr. nee Baird, Rept. 1853 pe^dejndella. 

C'hipeta Hulst, Can, Ent. xxiv. p. 62 (1892) perlepidella. 

Proboscis aborted and minute ; palpi downcurved, extending 
about twice the length of head and roughly scaled ; rnaxillai'v 
palpi well developed, filiform ; frons smooth and rounded ; an- 
tennfe of male minutely ciliated, the shaft slightly curved at base. 
Fore wing narrow, the apex rounded ; vein 2 fi'om towards angle 
of cell ; 3 and 5 from a point or separate ; 6 from below upper 
angle : 9 absent ; 10, 11 fi'om cell. Hind wing with vein 2 from 
just before angle of cell; 3, 5 strongly stalked, 4 absent; 6, 7 
from upper angle ; 8 anastomosing strongly with 7. 

(1) Bandera binotella. 

Anerastia binotella Zell. Yerh. zool.-bot. Ges. Wien, 1872, p. 108 ; 



90 SIR GEORGE HAMPSON ON THE 

Rag. Rom. Mem. viii. p. 409, pi. 44. f. 23; Dyar, Cat. Lep. 
N, Am. p. 440. 

U.S.A., Texas, New Mexico. 

(2) *Bandera perlepidella. 

Ghipeta jjerlepidella Hulst, Can. Ent. xxiv. p. 62 (1892); Dyar, 
Cat. Lep. N. Am. p. 441. 
U.S.A., Florida. 

(3) Bandera cupidinella. 

Bandera cupidinella Hulst, Ent. Am. iv.p. 119 (1888); Rag. 
Rom. Mem. viii. p. 410, pi. 45. f. 3 ; Dyar, Cat. Lep. iST. Am. 
p. 440. 

U.S.A., Colorado, New Mexico. 

(4) *Bandera subluteella. 

Bandera suhluteella Rag. IST. Am. Ph3^c. p. 19 (1887) ; id. Rom. 
Mem. viii. p. 410, pi. 40. f. 21 ; Dyar, Cat. Lep. N. Am. p. 440. 
U.S.A., Colorado, California, ISTew Mexico. 

AvjCtoruini. 

Bandera virginella Dyar, Proc. Ent. Soc. Wash. x. p. 116 

(1908) U.S.A., Washington. 

,, cameella Barnes & McD. Contr. ISTat. Hist. Lep. 
N.Am. ii. 4. p. 184, pi. i. f. 5 (1913^ 

U.S.A., Florida. 

Genus Laurentia. 

Type. 

Laurentia Rag. Nouv. Gen. p. 49 (1888) inclarella. 

Proboscis absent ; palf)i of male upturned to vei'tex of head, 
thickly scaled and hollowed out to receive the brush-like maxillary 
palpi, of female long and downcurved ; frons smooth, with tuft 
of scales ; antennae of male ciliated, the shaft with sinus at base 
of shaft containing a ridge of scales. Fore wing narrow, the 
apex rounded, tlie termen obliquely curved ; vein 2 from towards 
angle of cell ; 3 and 5 from angle, 4 absent ; 6 from below upper 
angle; 8, 9 stalked; 10, 11 from cell. Hind wing with vein 2 
from before angle of cell ; 3 and 5 from angle, closely approximated 
towards base, 4 absent; €, 7 shortly stalked; 8 anastomosing 
with 7. 

(1) tLAURENTIA ALBIVENELLA, Sp. n. 

S . Head and thorax creamy white mixed with some red- 
brown ; abdomen white, dorsally tinged with fulvous yellow 
towards base ; legs tinged with brown. Fore wing creamy 
white irrorated with some dark brown, the veins white deiined 



PYRALID^, SUBFAMILY HYPSOTROPIN/E. 91 

on each side by fine dark brown streaks, the median nervure 
more strongly streaked with white; a dark browi^ shade through 
upper part of cell and thence to termen below apex ; the inter- 
spaces between veins 4 and 2 shaded with .brown; cilia chequered 
with brown except at base. Hind wing white, the costal area 
and termen tinged with brown. 

Formosa, Takow (Wileman), 1 c? type. Uxp. 22 mm, 

• (2) *Laurentta inclarella. 

Laurentia inclarella Rag. Nouv. Gen. p. 49 (1888) ; id. Rom. 
Mem. viii. p. 408, pi. 38. f. 14. 

Java. 

Gen VIS Calamotropa, no v. 

Type, C pulverivena. 

Proboscis absent ; palpi downcurved, about two and a half 
times length of head and thickly scaled ; maxillary palpi well 
■ developed and slightly dilated with scales ; frons Avith large 
pointed conical prominence clothed with rough scales ; antennfe 
of female minutely ciliated. Fore wing long and narrow, the 
apex rounded ; vein 2 from towards angle of cell ; 3 and 5" 
sepai-ate, 4 absent; 6 from below upper angle ; 8, 9 stalked ; 10, 
11 from cell. Hind wing with vein 2 from just below angle of 
cell ; 3 and 4 absent ; 6, 7 shortly stalked ; 8 not anastomosing 
with 7. ' 

tCALAMOTROPA PULVERIVENA, Sp. n. 

$. Head and thorax ochreous slightly tinged with brown; 
pectus, legs, and abdomen ochreous white. Fore wing ochreous 
with diffused fuscous irroration along the veins except on costal 
area; a slight black point at lower angle of cell. Hind wing 
ochreous white, the costal area and termen tinged with brown. 

W. Australia, Sherlock R. {Clements), 3 5 type. Exp. 
24 mm. 

Genus Epidauria. 

Type. 
Epidauria Rag. Rom. Mem. viii. p. 405 

( 1901) transversariella. 

Proboscis absent ; ]Dalpi downcurved, extending' about three 
times length of head and thickly scaled ; maxillary palpi filiform ; 
frons smooth, with tuft of scales ; antennae of male laminate. 
Fore wing narrow, the apex lounded, the termen evenly curved; 
vein 2 from towards angle of cell ; 4 absent ; 5 from above angle ; 6 
from below upper angle; 8, 9 stalked ; 10, 11 from cell. Hind 
wing with vein 2 from before angle of cell ; 3 and 5 from angle ; 
4 absent ; 6, 7 from upper angle ; 8 approximated to but not 
anastomosing with 7. 



92 ' SIR GEORGE IIAMPSON ON THE 

(1) fEpiDAURIA PKRFASCIELLA, Sp. n. 

2 . Head and thorax whitish suffused with rufous ; abdomen 
whitish suffused with rufous and with obscure dark dorsal seg- 
mental bands. Fore wing whitish suffused with pale purplish 
red and slightly irrorated with blackish, the costal area rather 
whiter; a rather diffused black fascia from base through upper 
p.xrt of cell to termen below apex, forking in end of cell on vein 5 
to well beyond the cell ; vein 1 white at middle with rather 
diffused short black streaks above and below it ; the postmedial' 
line represented by a slight black mark above vein 6, a very 
oblique line between veins 5 and 2 and a short streak above 
vein 1 ; a terminal series of minute rather V-shaped black marks, 
Hiiid wing whitish saft'used with redilish brown ; cilia oclireous 
white with a slight brown line near base. 

Sierra Leoxe [Clements), 1 $ type. Exp. 28 mm. 

(2) *Epidauria tkans-versariella. 

Anerastia transversariella Zell. Isis, 1848, p. 588 ; Herr.-Schiiff. 
Enr. Schraett.iv. p. 109. Tin. f. 33 ; Rag. Rom. Mem. viii. p. 406, 
pi. 38. f. 17 ; Stand. Cat. Lep. pal. ii. p. 12, 

Dalmatia ; Corfu ; Asia Minor. 

(3) tepidauria chionocraspis, sp. b. 

2 . Head white tinged with purplish red ; thorax pale purplish 
red ; -pectus, legs, and abdomen white tinged with brown. Fore 
wing with narrow silveiy white costal fascia, the area below it 
dark chocolate-brown to median nervure and on terminal area 
dift'used on its lower side ; the rest of wing pale purplish red. 
Hind wing white with a slight ochreous tinge, the costa tinged 
with brown. 

Br. C. Africa, Mt. Mlanje (iVeave), 1 $ type. Exp. 24 mm. 

(4) Epjdauria phceniceella. 

Epidaurla pJiceniceella Rag. Bull. Soc. Ent. Fr. 1895, p. ciii ; 
id. Rom. Mem, viii. p. 407, pi. 52. f, 5 ; Stand. Cat, Lep. pal. ii, 
p. 12.- 

Asia Minor; Syria. 

(5) *Epidauria discella. 

Epidanrm discella Rag. Rom, Mem. ii. p. 407, pi. 40. f. 14 
(1901); Stand. Cat. Lep. pal. ii. p. 12. 
Mesopotamia- 

(6) *Epidauria strigosa. 

Anerastia strigosa Stand. Hor. Soc. Ent. Ross. xv. p. 225 
(1879); Rag. Rom. Mem. viii. p. 407, pi. 45. f. 2 ;' Stand. Cat. 
Lep. pal. ii. p. 12. 

Dalmatia ; Asia Minor ; Syria ; E. Siberia. 



PYRALID^, SUBFAMILY HYPSOXROPINiE. 93 

(7) tEPIDAUEIA SUBCOSTELLA, Sp. 11. 

5 . Head, thorax, and abdomen wliitish mixed with brown, 
the last dorsally fulvous yellow towards base; pectus, legs, and 
ventral surface of abdomen whitish suffused with dark brown. 
Fore wing whitish suffused with ochreous brown, the costa with 
a white streak below it to beyond middle defined below by a dark 
brown streak extending to apex. Hind wing whitish su£ui?ed 
with brown, the inner area whiter. 

Yunnan, Teng. Yeuk (Ilohson), 1 $ type. Ux]}. 26 mm. 

Genus Saluria. 

Type. 
Sahiria Rag. Ann. Soc. Ent. Fr. 1887, p. 258. maculivittelhi. 

Poujadia Kag. Nouv. Gen. p. 42 (1888) sepicosteUa. 

' Baroda Rag. Nouv. Gen. p. 42 (1888) paucigrcqihella. 

Goya Rag. JSTouv, Gen. p. 43 (1888) albivenella. 

Feclinigeria Rag. Nouv. Gen. p. 43 (1888, 

Apr.) macrella. 

Cayvga Hulst, Ent. Am. iv. p. 116 (1888, 

tSept.) geminatella . 

Atascosa Hulst, Trans. Am. Ent. Soc. xvii. 

p. 210(1890) glareosella. 

T'arramalta Hmpsn. Rom. Mem. viii. p. 366 

(1901) ensiferella. 

Ollia Dyar, J. N. Y. Ent. Soc. xii. p. 107 

(1904) santaritella. 

E%m%oorea Dyar, Insec. Inscit. Meustr. v. p. 91 

(1917) „ anchridis. 

Proboscis aborted and minute; palpi of male typicall}^ down- 
curved, extending about three times length of head and thicLly 
scaled, the 3rd joint moderate; maxillary palpi filiform; frons 
smooth with slight tuft of hair; antenna? of male typically with 
long uniseriate branches and without sinus and ridge of scales at 
base of shaft. Fore wing long and narrow, the apex rounded, 
the term en evenly curved ; vein 3 from close to angle of cell ; 
4, 5 strongly stalked ; 6 from below npper angle ; 8, 9 stalked ; 
10, 11 from cell. Hind wing with vein 2 from close to angle of 
cell ; 3 and 5 strongly stalked, 4 absent ; 6, 7 shortly stalked ; 
8 typically not anastomosing with 7. 

Sect. I. Palpi of male hollowed out to receive tlie brush-like maxillary palpi. 
A. AnteuniE of male with a sinus at base of shaft containing a ridge of scales. 
a. Antenna3 of male pectinate with long uniseriate branches. 
a'. Palpi of male oblique. 

(1) Saluria erodella. 

Poujadia erodella Rag. Nouv. Gen. p. 48 (1888); id. Rom. 
Mem. viii. p. 347, pi. 37. f. 5; Hmpsn. Moths Ind. iv. p. 59. 
Poujadia par vivluinella Hmpsn. Moths Ind. iv. p. 59 (1896). 
Punjab ; Bengal ; Madras ; Ceylon. 



94 SIR GEORGE HAMPSON ON THE 

6'. [Baroda). Palpi of male downcurved. 

(2) Salukia ctenucha. 

Poujadia ctemccha Turnei-, Pr. E,. Soc. Queensl. xxiv. p. 118 
(1913). 

N. Australia. 

(3) *Saluria flammella. 

Baroda Jlammella Hnipsn. Rom. Mem. viii. p. 348, pi. 51. f. 12 
(1901). 
Gambia. 

(4) fSALURIA CAKNESCENS, Sp. n. 

Head and thorax pale rufous ; abdomen white tinged with 
rufous, dorsally fulvous yellow towards base ; pectus and legs 
whitish suffused with rufous. Fore wing with the inner half pale 
purplish pink shading to rufous towards the white costal fascia, 
narrowing to apex ; the costal edge pale purplish red on basal 
half. Hind wing white with an ochreous tinge. 

Argentina, Sta. Fe, Ocampo [Wagner), 1 cf type, Gran Ohaco, 
Floi-enzia ( Wagner), J. $ . Exj). 28-30 mm. 

(5) fSALURIA PSAMMAIffiLLA, Sp. 11. 

c? . Head, thorax, and abdomen white tinged with ochreous 
brown ; antenn;© red-brown, white towards base. Fore wing 
white tinged with ochreous brown, the costal area rather whiter. 
Hind wing white faintly tinged with ochreous on costal half ; a 
fine dark terminal line. 

$ . Fore wing more strongly suffused with ochreous bi'own 
leaving a distinct white costal fascia narrowing to apex. 

Algeria, Hammam-es-Salahin {Walsingham), 1^,1$ type. 
Exp. 26-32 mm. 

(6) fSALURIA PAUOIGRAPHELLA. 

Barod,a faiicigraphella Rag. Nouv. Gen. p. 48 (1888); id. Rom. 
Mem. viii. p. 348, pi. 38. f . 5 ; Hmpsn. Moths Ind. iv. p. 60. 
Bengal, Calcutta ; Bombay, Kutch ; Madras. 

6. Antennse of male pectinate with short uniseriate branches; palpi oblique. 

(7) fSALURIA HEMIPH^ALIS. 

Saluria hemi2)hcealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi. 
p. 1251 (1912). 
Ceylon. 

(8) fSALURIA semirosella, sp. n. 

Head whitish suffused with rufous ; thorax pale pui-plish pink 



PYRALIDiE, SUBFAMILY HYPSOTROPINiE. 95 

with some rufous on. shoulders ; abdomen white tinged with red- 
brown, dorsally fulvous yellow towards base ; pectus and legs 
white tinged with red-brown. Fore wing with the inner half 
pale purplish pink, the veins finely streaked with white; the 
cell and area beyond it red-brown below the white costal fascia 
narrowing to apex ; the costal edge red-brown towards base, then 
pale pink to beyond middle. Hind wing white tinged with red- 
brown, the cilia whiter with a red-brown line near base. 

Sierra Leone {Clements), 1 c? , 2 $ ; Uganda, Nakwai Hills 
{Lowe), 1 J, Gondokoro {Reymes-Cole), 1 S type. Exp. 16- 
20 ram. 

c. Antennae of male strongly serrate ; palpi downcurved, 

(9) fSALURIA MINUTELLA. 

Saluria ininutella Hmpsn. J. Bomb. Nat. Hist. Soc. xv.,. p, 20 
(1903). 

Bombay ; Ceylon, sp. n. 

(10) tSALURIA NILGIRIENSIS, Sp. n. 

(S . Head and thorax pale rufous ; abdomen white tinged with 
rufous ; antennae with some black in the ridge of scales ; pectus, 
legs, and ventral surface of abdomen brown. Fore wing with the 
inner half creamy white suffused with rufous varying to purplisli 
pink ; the upper part of cell and area beyond it brown below the 
white costal fascia narrowing to apex ; the costal edge brown 
towards base, then irrorated with rufous ; a rather oblique post- 
medial series of shoi't dark streaks on veins 6 to 1 and a terminal 
series of slight dark points. Hind wing white tinged with brown 
especially on costal area, the cilia whiter with a slight brown line 
near base. 

Madras, Nilgiris, Pj'^kara {Andrewes), 4 S type. Exjj. 26- 
28 mm. 

d. (Poiijadia). Antennae of male laminate ; palpi oblique. 

(11) Saluria SEPICOSTELLA. 

Poujadia sepicostella Rag, Nouv. Gen. p. 42 (1888); id. Rom. 
Mem. viii. p. .345, pi. 36. f . 2 ; Hmpsn. Moths Ind. iv. p. 58. 
Formosa ; Punjab ; Borneo, Pulo Laut. 

(12) Saluria claricostella. 

Poujadia claricostella Rag. Nouv. Gen. p. 42 (1888) ; id. Rom. 
Mem. viii. p. 345, pi. 36. f. 26. 
Gold Coast ; Br. C. Africa. 

(13) tSALURiA GLAREOSELLA. 

Anerastia glareosella Zell. Yerh. zool.-bot, Ges. Wien, 1872, 



96 SIR GEORGE HAMPSON ON THE 

p. 553 ; Rag. Rom. Mem. viii. p. 345, pi. 37. f. 19 ; Dyar, Cat. 
Lep. ]Sr. Am. p. 438, 
U.S.A., Texas. 

(14) fSALURIA STICTOPHOEA, Sp. n. 

Head and thorax reddish brown ; abdomen whitish tinged 
with brown, dorsally fulvous yellow towards base ; antennae of 
male with the ridge of scales black : pectus, legs, and ventral 
surface of abdomen white mixed with brown. Fore wing whitish 
suffused with brown and thickly irrorated with dark brown ; a 
white costal fascia slightly irrorated with dark brown and rather 
diffused below; a small round black discoidal spot; an indistinct 
rather diffused blackish postmedial line not quite I'eaching the 
costa ; cilia with a fine white line at base. Hind wing whitish 
suffused with brown. 

Mashonaland, Salisbury (Marshall), 1 J , 1 $ type. Exj[). 
20 mm. 

(15) Saluria spurcella. 

Poujadia spui^cella Rag. ISTouv. Gen. p. 42 (1888) ; id. Rom. 
Mem. viii. p. 346, pi. 37. f. 4 ; Hmpsn. Moths Ind, iv. p. 59. 
Punjab ; Bombay ; Madeas ; Burma ; Labuan. 

(16) fSALUEIA INFICITA. 

Acrohasis inficita Wlk. xxvii. 30 (1863); Hmpsn. Moths Ind. 
iv. p. 58 ; Rag. Rom. Mem. viii. p. 346, pi. 44. f. 18. 
Teansvaal ; Madeas ; Ceylon. 

(17) tSALUEIA FLAVICOSTA, sp. n. 

Head rufous ; thorax grey-brown, the tegulse dorsally rufous ; 
abdomen grey-brown, the 2nd and 3rd segments dorsally fulvous 
yellow, the anal segment pale yellow ; antennae brown ; palpi 
yellow with black patch on 2nd segment above ; pectus, legs, and 
ventral surface of abdomen grey-brown and whitish. Fore wing- 
dark brown glossed with grey ; a narrow pale yellow costal fascia, 
the costal edge dark towards base. Hind wing pale grey-brown, 
the cilia whitish with a brown line near base. 

SiEEEA Leone {Clements), 1 c5' ; N. Nigeria, Zungeru {Macfie), 
1 (S type. Exj). 16 mm. 

(18) *Salueia floscella. 

Atacoea Jioscella Hulst, Trans. Am. Ent. Soe. xvii. p. 210 
(1890); Rag. Rom. Mem. viii. p. 346, pi. 51. f. 4; Dyar, Cat. 
Lep. K Am. p. 438. 

U.S.A., Texas. 

(19) tSALUEIA STICTELLA, Sp. n. 

S . Head and thorax brownish ochreous ; abdomen ochreous 



PYRALID.E, SUBFAMILY HYPSOTROPINiE. 97 

white, dorsally fulvous yellow towards base ; palpi white below 
towards base ; pectus, legs, and ventral surface of abdomen 
white, the legs tinged with brown. Fore wing white tinged with 
ochreous brown except the costal area to discal fold, irrorated 
with black scales, the costal and inner areas less irrorated ; small 
obliquely placed black antemedial spots in and below the cell and 
a discoidal spot ; a postmedial series of black points on veins 6 
to 1 and a terminal series of slight black points. Hind wing- 
white tinged with ochreous brown. 

Bahamas, Andros (J^onhote), 1 c? type. Bxp. 16 mm. 

(20) Saluria ochridorsella. 

Poujadia ochridorsella Rag-. Nouv. Gen. p. 42 (1888); id. Rom. 
Mem. viii. p. 347, p. 37. f. 1 ; Hmpsn. Moths Ind. iv. p. 58. 
Punjab ; Bengal ; Ceylon. 

/21) t)SALURIA ROSELLA. 

Poujadia rosella Hmpsn. Moths Ind. iv. p. 59 (1896); id. Rom. 
Mem. viii. p. 347, pi. 51. f. 20. 
Madras, Nilgiris. 

(22) fSALURIA VERECUNDELLA. 

Poujadia verecunddla Rag. Rom. Mem. viii. p. 347, pi. 37. f. 2 
(1901). 
Colombia ; S. Brazil ; Argentina. 

(23) fSALURIA LENTISTRIGELLA, Sp. n. 

Head and thorax white, the tegula? except dorsally and the 
pa,tagia slightly tinged with rufous ; abdomen white slightly 
tinged with rufous, dortally fulvous yellow towards base. Fore 
wing chalky white, the veins defined by slight streaks of red- 
bi'own scales and the inner area slightly irrorated with red- 
brown ; a faint rufous shade below median nervure; a small 
rather difi'used round blackish spot on vein 1 and slight obliquely 
placed postmedial spots on veins 2 and 1 ; a fine punctiform 
blackish terminal line ; cilia chequered with blackish near base 
except towards torn us. Hind wing white with fine bi-own ter- 
minal line except towards tornus. 

Sierra Leone {Clements), 1 S ; Ctold Coast. Bibianaha 
{Spiirrell), 2 cJ , 1 $ type. Pxp. 22-24 mm. 

(24) tSALURIA desertella, sp. n. 

2 ■ Head and thorax white faintly tinged with rufous ; 
abdomen white, dorsally fulvous yellow towards base. Fore 
wing white tinged with ochreovis, the cilia whiter. Hind wing- 
white. 

N. Australia, Alexandria (Stalkei-), 3 5 ; W. Australia, 
Sherleck R. {Clements), 1 $ type. A'a-jw. 22-32 mm. 

Proc. Zool. Soc— 1918, No. VII. 7 



&8 SIR GEORGE HAMPSON ON THE 

B. (Eumoorea). Anteimaj of male laminate, witliout sinus and ridge of scales 
at base of shaft. 

(25) *Saluria anchribis. 

Eumoorea anchridis Dyar, Insec. Iiiscit. Meutv. v. p. 91 (1917). 
Br. Guiana. 

(26) fSALURIA flavip.urpurella, sp. n. 

(S . Head and thorax rufous mixed with ochreous ; abdomen 
ochreous rufous, dorsally fulvous yellow towards base ; pectus 
and legs white suftused with brown ; venti-al surface of abdomen 
brown, whitish at base. Fore wing with creamy white costal 
fascia narrowing to apex, the costal edge blackish at base, defined 
below by dark brown followed by deep piirple-pink to just below 
the cell and vein 4, the inner half of wing yellow. Hind wing 
ochreous white, the apical area tinged with brown. 

Paraguay, Sapucay (Foster), 1 c? type. U.v]). 16 mm. 

Sect. II. Maxillar}' palpi of male filiform. 

A. Antennae of male with a sinus at base of shaft containing a ridge of scales. 
a. Antennae of male pectinate with rather long uniseriate branches to about 
half length ; palpi downcurved. 

(27) tSALURIA PARANENSIS. 

Pectinigeria paranensis Hmpsn. Rom. Mem. viii. p. 355, pi. 55. 
f. 6 (1901). 
S. Brazil. 

(28) *Saluria mus^ella. 

Pectinigeria musceella Schaus, A. M. N. H. (8) xi. p. 239 (1913). 
Costa Rica. « 

(29) fSALURIA TENUICOSTA, Sp. n. 

Head and thorax rufous tinged with pale pink ; abdomen 
ochreous, dorsally fulvous yellow towards base ; pectus, legs, and 
ventral surface of abdomen whitish suffused with red- brown. 
Fore wing pale purplish red sparsely irrorated with black ; a 
narrow white costal fascia defined below by blackish to beyond 
middle ; a terminal series of slight dark points. Hind wing 
white, tinged with ochreous bi'own on costal area and termen, 

Paraguay, Sapucay (Foster), 1 (5 , 1 $ type. Fxp. 18 mm. 

(30) Saluria pleubostioha. 

Ilypsotropha pleurosticha Turner, Pr. R. Soc. Qneensl. xviii. 
p. 115(1903). 
Queensland. 

(31) fSALURIA neuricella, sp. n. 

Head and thox^ax rufous mixed with some Avhitish ; abdomen 



PYRAIilD^, SUBFAMILY HYP80TROPIN^. 99 

ochreous white, dorsally fulvous yellow towards base; pectus and 
legs white sufl'used with brown. Fore wing rufous tinged with 
purplish pink, the veins streaked with white. Hind wing whitish 
suffused with ochreous brown, the cilia whiter. 

Queensland, Peak Downs, 1 cJ, 1 ? type, Exp. 22 mm. 

h. Antennae of male laminate and ciliated. 
a'. {Goya). Palpi oblique, the 3vdjoint porrect. 

(32) Saluria albivenella. 

Goya alhivenella Rag. Nouv. Gen. p. 4'i (1888) ; id. Rom. Mem. 
viii. p. 349, pi. 37. f. 6. 
S. Brazil; Argentina. 

(33) *Saluria varicosella. 

Goya varicosella Rag. Nouv. Gen. p. 43 (1888); id. Rom. Mem. 
viii: p. 350, pi. 37. f. 7. 
Gambia. 

(34) fSALURIA proleucella, sp. n. 

S . Head and thorax white suffused with red-brown ; abdomen 
white tinged with red-brown, dorsally fulvous yellow towards 
base. Fore wing white tinged and irrorated with i-ed-brown, the 
veins finely streaked with white ; a white costal fascia slightly 
irrorated with red-brown, narrowing to apex. Hind wing creamy 
white, the costal area faintly tinged with brown. 

Gold Coast, Accra [Sir G. Carter), 1 c? type. Exp. 22 mm. 

(35) tSALURIA grammivena, sp. n. 

Head, thorax, and abdomen white suffused with red-brown ; 
pectus, legs, and ventral surface of abdomen white irrorated with 
red-bi'own. Fore wing white thickly irrorated with red-bi'own, 
the veins strongly streaked with white and defined on each side 
by fine red-brown streaks ; cilia whitish. Hind wing white 
tinged with red-brown. 

N". Australia, Alexandria {Stalker), 1 J ; W. Australia, 
Sherlock R. (Clements), 1 $ type. Exp. 26 mm. 

b'. (Fectinigeria). Palpi downcurved. 

(36) Saluria nigritella. 

Pectinigeria nigritella Rag. Rom. M^m. viii. p. 353, pi. 37. f. 13 
(1901). 

Gaboon ; N. Rhodesia. 

(37) *Saluria ardiperella. 

Altoona ardiferella Hulst, Ent. Am. iv. pi. 118 (1888); Rag, 
Rom. Mem. viii. p. 506, p. 24. f. 14 : Dyar, Cat. Lep. N. Am. 
p. 439. 

7* 



100 SIR GEORGE HAMPSON ON I'HE 

Aurora nigromaculella Hulst, J. N. Y. Ent. See. viii. p. 224 
(1900) ; Dyar, Oat. Lep. N. Am. p. 438. 

U.S.A., Texas, Colorado, California, N. Mexico, Arizo)ia. 

(38) SaLURIA DEVyLDERI. 

Pectinigeria devylderi Rag. Noiiv. Gen. p. 43 (1888); id. Rom. 
Mem. viii. p. 354," pi. 37. f. 10. 

N. Rhodesia: Damaraland ; Natal. 

(39) Saluria arcticostella. 

Pectinigeria arcticostella Rag. JSTouv. Gen. p. 43 (1888); id. 
Rom. Mem. viii. p. 3.54, pi. 37. f. 9. 
N. Nigeria ; Zanzibar. 

(40) *Saluria furvella. 

Pectinigeria furvella Rag. Nouv. Gen. p. 43 (1888) ; id. Rom. 
Mem. viii. p. 355, pi. 37. f. 11. 
Madagascar. 

(41) Saluria macrella. ' 

Pectinigeria macrella Rag, ISTouv. Gen. p. 44 (1888) ; id. Rom. 
Mem. viii. p. 355, p]. 37. f. 16. 
Br. C. Africa ; JSTatal. 

(42) Saluria subcarnella. 

Pectinigeria snhcarnella Rag. Nouv. Gen. p. 44 (1888); id. 
Rom. Mem. viii. p. 356, pi. 37. f. 1.5. 

N. Nigeria ; Transvaal ; Natal ; Cape Colony. 

(43) Saluria gemmatella. 

Spermatophora geinmatella Hulst, Ent. Am. iii. p. 134 (1887); 
Rag. Rom. Mem. viii. p. 356, pi. 37. f. 12 ; Dyar, Cat. Lep. 
N. Am. p. 438. 

U.S.A., Illinois, Colorado, California. 

(44) fSALURIA RHODOPHiEA, Sp. n. 

§ . Head and thorax vinous purple ; abdomen fulvous tinged 
with brown except dorsally towards base ; palpi, pectus, legs, and 
ventral sui'face of abdomen I'ed-brown. Fore wing vinous purple 
with a slight whitish streak on subcostal nervure tending to 
fork at extremity and defined below by a dark fascia, extending 
through the cell to apex. Hintl wing ochreous white tinged witli 
brown especially towards costa. 

Transvaal (Pead), 1 $ type. Exp. 24 mm. 

(45) tSALURIA SUBCOSTELLA, sp. n. 

$ , Head and thorax deep rufous mixed with some whitish 
especially on metathorax ; abdomen fulvous with slight whitish 



PYRALIDiE, SUBFAMILY HYPSOXROPIN^. 101 

segmental lines and the extremity whitish ; pectus, legs, and 
ventral' surface of abdomen whitisli suflused with red-brown. 
Fore wing whitish tinged with red-brown; the costal edge dark 
brown to well beyond niiddle ; a white costal fascia narrowing to 
apex, defined below by a dark brown fascia. Hind wing ochreous 
white tinged with brown especially towards costa. 

Br. C. Africa, Mt. Mlanje {Reave), 1 $ type. Ex'p. 30 mm. 

(46) fSALURIA DISTICTELLA, sp. n. 

<S . Head and thorax whitish, the vei'tex of head, tegulaj and 
patagia sufi'used with red-brown ; antennae tinged with dark 
brown ; palpi with some blackish scales at side of 2nd joint ; 
abdomen white, dorsally fulvous yellow towards base ; legs tinged 
with red-brown. Fore wing white tinged with flesh-pink ; a 
pure white costal fascia narrowing to aj)ex and defined below by 
rufous which is rather difi'used below; an antemedial red-brown 
point on vein 1 and medial point in submedian fold. Hind wing 
white. 

Queensland, Brisbane, 1 J type Ex2). 20 mm. 

(47) tSALURIA rufella, sp. n. 

5 . Head and thorax rufous ; abdomen whitish suffused with 
brown, dorsally fulvous yellow towards base ; pectus and legs 
whitish suffused with rufous. Fore wing uniform rufous. Hind 
wing ochreous white. 

Bengal, Oudh [Pilcher), 1 5 type. Ex-ji. 26 mm. 

(48) tSALURIA OPIFICELLA. 

Anerastia opificella Zell. Stett. Ent. Zeit. 1867, p. 406 ; Hmpsn 
Moths Ind. iv. p. 60 ; Rag. Rom. Mem. viii. p. 357, pi. 44. f. 16. 
Kashmir ; Bengal ; Ceylon ; Burma. 

(49) *Saluria breviculella. 

Sahirla hreviculella Hmpsn. J. Bomb. Nat. Hist. Soc. xii. p. 308 
(1898); id. Rom. Mem. viii. p. 358, pi. 37. f. 14. 
Bombay. 

\^■ Anteunie of iiiiilo without sinus and ridge of scales at base of shaft. 

a. {Pari'amatta). Hind wine with vein 8 anastomosing with 7; antennte of 
male laminate and ciliated. 

(50) SaLURIA DICHROELLA. 

>Sakiria dichroella Rag. Ent. Am. v. p. 113 (1889); id. Rom. 
Mem. -ftiii. p. 363, pi. 39."f. 1 ; Dyar, Cat. Lep. N. Am. p. 439.- 
U.S.A., Texas. 

(51) *Saluria neotomella. 

Eiicarphia neotomella Meyr. Proc. Linn. Soc. N. S. W. iv. p. 226 
(1879) ; Rag. Rom. Mem. viii. p. 363, pi. 37. f 25. 
N. S. Wales. 



102 sir george hampson on the 

(52) *Sal,uria ostreblla. 

Sahiria osireella Rag. N. Am. Phyc. p. 18 (1887); id. Rom. 
iM6m. viii. p. 362, pi. 38. f. 6 ; Dyar, Oat. Lep. N. Am. p. 438. 
U.S.A., Arizona. 

(53) tSALURIA tetradblla. 

Anerastia tetradella Zell. Verb, zool.-bot. Ges. Wien, 1-872, 
p. 552 : Rag. Rom. Mem. viii. p. 362, pi. 42. f. 23 ; Dyar, Cat. 
Lep. N. Am. p. 439. 

U.S.A., Texas, Colorado. 

(54) Saluria holochroa. 

Poiojadia holochroa Turner, Pi-. R. Soc. Queensl. xviii. p. 121 
(190.3). 

W. Australia ; Victoria. 

(55) *Saluria cancelliella. 

Saluria cancelliella Rag. IS^ouv. Gen. p. 44 (1888); id. Rom. 
Mem. viii. p. 363, pi. 37. f. 23. 
Brazil. 

(56) *Saluria ensiferella. 

Uticarphia ensijiherella Meyr. Proc. Linn. Soc. N. S. W. iii. 
p. 209 (1878) ; Rag. Eom. Mem. viii. p. 366, pi. 39. f. 2. 
N. S. Wales. 

b. Hind wing with vein 8 not anastomosing with 7. 
a'. (Saluria). Antennie of male pectinate with long uniseriate branches. 

(57) *Saluria pectigerella. 

/Saluria pectigerella Rag. Ann. Soc. Eut. Fr. 1887, p. 259 ; id. 
Rom. Mem. viii. p. 360, pi. 37. f. 24 ; Stand. Cat. Lep. Pal. ii. 
p. 13. 

W. Turkestan. 

(58) Saluria maculivittella. 

Saluria maculivittella Rag. Ann. Soc. Ent. Fr. 1887, p. 258; 
id. Rom. Mem. viii. p. 361, pi. 38. f. 7; Stand. Cat. Lep. pal. 
ii. p. 13. 

Saluria armeniella Rag. Nouv. Gen. p. 44 (1888). 

Algeria; Tunis; Armenia; Cyprus; W.Turkestan; Ceylon. 

(59) tSALURIA pulverosa. 

Poujadia pulverosa Hmpsn. Moths Ind. iv. p. 60 (1896); id. 
Rom. Mem. viii. p. 361, pi. 37. f. 3. 

SiND. 



pyralid^, subfamily hypsotropin^e. 103 

(60) *Saluria magnesiella. 

Sahiria magnesiella Rag. Nouv. Gen. p. 44 (1888); id. Rom. 
Mem. viii. p. 360, pi. 37. f. 18 ; Stand. Oat. Lep, pal. ii p. 13 
W. Turkestan. 

b' . Autemiae of male serrate and ciliated. 

(61) tSALURIA MESOMBLANELLA, Sp. n. 

$ . Head purple-red and black-brown ; thorax purple-red and 
whitish ; abdomen whitish suffused with brown, dorsally rufous 
towards base ; pectus, legs, and ventral surface of abdomen 
whitish suftused with red-brown and dark brown. Fore wing 
whitish tinged with purplish pink, the costa and termen deep 
purple-crimson, the medial area suffused with black except at 
costa, and the costa defined below by black ; an antemedial black 
point on vein 1 with a purple-pink shade below it on inner 
margin ; the inner edge of the medial black area obliquely curved 
and its outer edge bent inwards below submedian fold ; cilia 
pale fuscous. Hind wing white tinged with brown especially at 
termen, the cilia with a fine brown line near base. 

Transvaal, "White R. {Cooke), 1 $ type. Exp. 28 mm. 

(62) Saluria tripartella. 

Saluria tripartella Rag. Rom. Mem. viii. p. 359, pi. 39. f. 19 
(1901). 

Transvaal; Natal; Basutoland. 

(63) Saluria callirhoda. 

Poujadia callh^hoda Turner, Pr. R. Soc. Queensl. xviii. p. 120 
(1903). 

Queensland. 

(64) tSALURIA inxbrpunctella, sp. n. 

Head, thorax, and abdomen ochreous suffvised with rufous; 
pectus, legs, and venti^al surface of abdomen reddish ochreous 
mixed with brown. Fore wing ochreous sufi'used with rufous ; 
the veins white defined on each side by fine black-brown streaks, 
the submedian fold and vein 1 defined by fine rufous streaks ; 
the costal edge black towards base ; an antemedial blaek point on 
vein 1, a point in lower angle of cell, an oblique postmedial series 
in the interspaces, incurved below submedian fold and a, terminal 
series. Hind wing ochreous with a tine biownish terminal line 
except towards tornus. 

Mashonaland (Bobbie), 2 d , 2 $ type ; Transvaal, Kranspruit 
[Janse), 1 5 , Pretoria {Distant), 1 $ , Exp. 20-24 mm. 

(65) tSALURIA INSIGNIFICELLA, Sp. n. 

$ . Head and thorax white mixed with red-brown ; abdomen 
white tinged with brown, dorsally fulvous yellow towards base. 



104 SIR GEORGE HAMPSON ON THE 

Fore wing whitish tinged with red-brown, especially towards 
costa; the costa narrowly white. Hind wing white tinged with 
red -brown. 

Paraguay, Sapucay (Foster), 1 5 type Exjj. 18 mm. 

Auctorum. 

Atascosa ijuadricolorella Dyar, Proc. Ent. 8oc. Wash. vi. p. 114 

( 1 904). Sect. Poujadia U.S. A. , N. Mexico. 

Poajadia piiuella Dyar, J. N. Y. Ent. Soc. xiv. p. 31 (1906). 

U.S.A., Arizona. 
Pectiitigeria 'pO'^ipoiierella Dyar, Proc. Ent. Soc. Wash. x. p. 107 

(1908) U.S.A., Colorado. 

Ollia parvella Dyar, J. N. Y. Eat. Soc. xiv. p. 31 (1906). Sect. 

Saluria U.S.A., Texas. 

Ollia honojyoiierella Dyar, Proc. Ent. Soc. Wash. x. p. 107 

(1 908). Sect. Saluria U.S.A., Arizona. 

Ollia santaritella Dyar, J. IST. Y. Ent. Soc. xii. pp. 107-8 (1904). 

Sect. Saluria U.S.A., Arizona. 

Poajadia leucoiieara Turner, Pr. R. Soc. Queensl. xxiv. p. 118 

(1913). ? Sect. Paramatta N. Australia. 

Pectiiiv/eria violodis Dyar, Pi-. U. S. Nat. Mus. xlvii. p. 347 

(1913) Panama. 

Poujadia cyttarella Dyar, Pi-. U. S. Nat. Mus. xlvii. p. 347 

(1913) ...'. Panama. 

Genus Prophtasia. 

Type. 

Prophtasia Rag. Ania. Soc. Ent. Fr. 1887, p. 259. . . platycerella. 

Proboscis aborted and minute ; palpi typically obliquely up- 
turned, tlie 3rd joint porrect ; maxillary palpi filiform; frous 
smooth, with slight tuft of scales ; antennae of male typically 
laminate, the shaft slightly carved at base and without distinct 
I'idge of scales. Fore wing long and narrow, the a]iex rounded, 
the termen evenly curved ; veins 2 and 3 typically from angle of 
cell ; 4, 5 stalked ; 6 from below upper angle ; 8, 9 stalked ; 10, 
1 1 from cell. Hind wing with vein 2 from well befoi'e angle 
of cell ; 3 and 5 from angle, typically appi'oxiniated for some 
distance, 4 absent ; 6, 7 shortly stalked ; 8 typically not anasto- 
mosing with 7. 

Sect. I. Fore wing witli veins 2, 3 from a point at angle of cell ; hind wing with 
vein 8 not anastoniossing with 7. 

(1) Prophtasia platycerella. 

Prophtasia platycerella Rag. Ann. Soc. Ent. Fr. 1887, p. 259 ; 
id. Rom. M6m. vi'ii. p. 252, pi. 37. f. 8 ; Stand. Cat. Lep. pal. 
ii. p. 13. 

Armenia. 



PYRALID^, SUBFAMILY H YPSOTROPIN^. 105 

Sect. II. Fore wiug with vein 2 from well before angle of cell, 3 from before angle ; 

hind wing with veins 3 and 5 not approximated towards base. 

A. Hind wing with vein 8 anastomosing with 7; palpi obliquely uptm'ned and 

hardly reaching to vertex of head, the maxillary palpi filiform ; antennae of 

male with the shaft slightly curved at base and without distinct ridge of 

scales. 

(2) fpROPHTASIA SPHALMATELLA, Sp. 11. 

c? . Hea'l and thorax black-brown tinged with grej' ; abdomen 
lirowiiish oclireous, dorsally t'nlvous yellow towards base, ventrally 
grey-bi'own. Fore wing black-brown mixed with some grey, 
especially on inner area ; a pure white fascia on costal area, 
leaving the costa bla,ck-l>rown and not reaching the apex ; an 
indistinct diffused dark antemedial spot on vein 1 and oblique 
postmedial line. Hind wing whitish suffused with brewn ; a. 
darker tern)iual line and white line at base of cilia. 

Mashoxaland, Salisbury {Marshall), 1 6 type. Exp. 28 mm. 

(3) fPROPHTASIA EPITEUXIS, Sp. n. 

S . Head and thorax glossy black-brown ; abdomen grey suf- 
fused with brown ; palpi white below towards base ; pectus, legs, 
and ventral surface of abdomen black-brown, the femora and 
tibiaB streaked with white. Fore wing glossy black-brown, the 
inner area tinged with grey ; a pure white costal fascia, narrow- 
ing to apex ; an indistinct diffused dark antemedial line from cell 
to inner margin and oblique postmedial line. Hind wing grey 
suffused with fuscous ; a slight punctiform dark terminal line and 
white line at baseof cilia. 

Burma, Hsipaw {de Niceville), 2 J type. Exi). 24-28 mm. 

(4) fPROPHTASIA GLAUCOPH^A, Sp. n. 

Head and thorax grey-brown tinged with white ; abdomen 
white suffused with grey-brown ; palpi white in front except 
towards tips ; pectus, legs, and ventral surface of abdomen white 
mixed with some brown. Fore wing pale grey-brown, black- 
brown towards the I'ather broad white costal fascia which does 
not narrow to apex. Hind wing creamy white slightly tinged 
with brown. 

Kashmir {Pilcher), 1 J ; Punjab, Kangra [Dudgeon), 1 J , 
Moghal iSarai (Betton), 1 cJ ; Bengal, Oudh [Filcher), 1 r? ; Sind, 
Karachi {Swinhoe), 1$; Madras, Belgaum [Watson), \ S type. 
Exj). 18-22 mm. 

(5) fPROPHTASIA AMPHICHEA, Sp. n. 

Head and thorax grey-brown mixed with white ; abdomen 
white tinged with brown ; pectus and legs white mixed with 
brown. Fore wing pale pur2:)lish brown mixed with wdiite, the 
costal area broadly white with slight bi'own irroration ; indistinct 
diffused dark curved ante- and postmedial lines except towards 



106 SIR GEOHGE UAMPSON ON THE 

cosba ; two slight blackish discoidal spots. Hind wing white 
tinged with brown ; a sHght dark spot at upper angle of cell and 
fine terminal line ; cilia whiter. 

Sierra Leone {Clements), 1 cT , 4 $ ; N. Nigeria, Minna {Macfie), 
1 6 type, Zungeru {Simpson), 1 5 • Exjj. 16-20 mm. 

B, Hind wing with vein 8 not anastomosing with 7. 
a. Palpi of male obliquely upturned to above vertex of head and hollowed out 
to receive the brush-like maxillary palpi, the 3rd joint porrect ; antennae 
serrate with sinus at base of shaft containing a large ridge of scales. 

(6) fPROPHTASIA pyrostrota, sp. n. 

(S . Head and thorax creamy white snfFused with rufous ; 
abdomen, pectus, and legs creamy white. Fore wing ochreous 
white iri'orated with fiery red, the costal fascia only defined by a 
red-brown fascia below it from base to apex ; minute nntemeclial 
black streaks on median nervure and veiit 1 and an obliijue post- 
medial series ; two slight black discoidal spots ; a tei-minal series 
of black points. Hind wing creamy white. 

Punjab, Kangra Valley, 4500' {Dudgeon), 1 S type. Exj). 
26 mm. 

h. Palpi of female downcurved and extending about three times length of head. 

(7) Prophtasia BISTRIATELLA. 

Cayuga histriatella Hulst, Trans. Am. Ent. Soc. xvii. p. 209 
(1890) ; Dyar, Cat. Lep. N. Am. p. 438. 

Peoria discostrigella Dyar, Proc. Ent. Soc. Wash. vi. p. 115 
(1904). 

U.S.A., N. Mexico, Arizona, 

Genus Aurora. 

Type. 
Aurora Rag. N. Am. Pliyc. p. 18 (1887) lonpipalpalla. 

Proboscis aborted and minute; palpi porrect, extending about 
four times length of head, thickly fringed with hair, the 2nd 
joint slightly downcurved ; maxillary palpi slight and filiform ; 
frons smooth, with long pointed tuft of hair; autennfe of female 
minutely ciliated. Fore wing long and narrow, the apex rounded, 
tlie termen obli(]uely curved ; vein .8 from close to angle of cell ; 
4, 5 strongly stalked ; 6 from below upper angle ; 8, 9, 10 stalked ; 
11 from cell. Hind wing, with vein 2 from close to angle of cell ; 
3 and 5 stalked, 4 absent ; 6, 7 shortly stalked ; 8 anastomosing 
with 7. 

t Aurora longipalpella. 

Aurora longijxdpeUa Rag. N. Am. Phyc. p. 18 (1887) ; id. Rom. 
Mem. viii. p. 337, pi. 44. f. 2 ; Dyar, Cat. Lep. N. Am. p. 437. 

U.S.A. 



PYRALID^, SUBFAMILY HYPSOTROPIN^. 107 

Genus Fossifroktia. 

TjT)e. 
Fossifrontia Hmpsn. Rom. Mem. viii. p. 388 

(1 901 ) leicGoneurella. 

Proboscis absent ; palpi upturned to about middle of frons, 
thickly scaled a.nd hollowed out to receive the brush-like maxillary- 
palpi ; frons with truncate conical prominence hollowed out in 
front ; antennae of male uniserrate and ciliated, the shaft with 
large sinus and ridge of scales at base ; fore femora with tuft of 
hair above, the mid tibia? fringed with long hair on outer side. 
Fore wing narrow, the apex rounded, the termen evenly curved ; 
vein 3 froin before angle of cell ; 4, 5 stalked ; 6 from below 
upper angle ; 8, 9, 10 stalked ; 11 from cell ; the male with small 
fold on underside from medial part of costa fringed with large 
scales and with some androconia below it above, in and below 
the cell. Hind wing with vein 2 froin towards angle of cell ; 

3 and 5 from angle and closely approximated for some distance, 

4 absent ; 6, 7 shortly stalked ; 8 closely approximated to 7 but 
not anastomosing with it. 

Fossifrontia leuconeurella. 

Fossifro7itia leuconeurella Hmpsn. Rom. Mem. viii. p. 339, 
pi. 52. f. 19 (1901). 

QuEEisrsLAND, Cooktown, Cedar Bay. 

Genus Commotria. 

^ . , . Type. 

Commotria Berg, An. Soc. Arg, xix. p. 278 

(1 885) i invenustella. 

Mangcda Rag. Nouv. Gen. p. 41 (1888) crassiscapella. 

Tolima Rag. Nouv. Gen. p. 41 (1888) oherthuri. 

Altoona Hulst, Ent. Am. iv. p. 116 (1888) ... opacella. 
Volusia Hulst, Trans. Am. Ent. Soc. xvii. 

p. 206 (1890), nee Rob. Desv. Dipt. 1830 ... roseopeiinella. ' 
Trivolusia Dyar, Cat. Lep. N. Am. p. 438 

(1 902) roseopemiella. 

Proboscis absent ; palpi downcurved, extending about three 
times length of head and thickly scaled ; maxillary palpi slight 
and filiform ; frons rounded and with short tuft of hair ; antenna? 
of male typically unipectinate, the apical part ciliated, the shaft 
with ridge of scales above at base. Fore wing long and nari'ow, 
the apex rounded, the termen obliquely curved ; vein 3 from 
angle of cell ; 4, 5 stalked ; 6 from below upper angle ; 8, 9, 10 
stalked; 11 from cell. Hind wing with vein 2 from angle of 
cell ; 3 and 5 strongly stalked, 4 absent ; 6, 7 shortly stalked ; 
8 anastomosing with 7. 



108 SIR GEORGE UAMPSON CUV THE 

Sect. I. (Commotria.) Antenna? of male unipectinate, the apical part ciliated. 

(1) Commotria mesiella, sp. u. 

Head and thorax purple-pitik mixed with some whiti.sh ; abdo- 
men pale fulvous yellow, ochreous white at base and extremity; 
pectus, legs, and ventral surface of abdomen whitish tinged with 
purple. Fore wing' purple-pink, the veins white defined on each 
side by fine brown streaks, the white on median nervui'e stronger 
and more strongly defined by blRck-brown below; a minute black- 
brown spot at upper angle of cell and point at lower angle. Hind 
wing whitish tinged with ochreou.s brown. 

Br. C. Africa, Mt. Mlanje (Neave), 9^,3$ type. iJxp. 
20-22 mm. 

(2) Commotria laticostella. 

Commotria laticostella Hmpsn. Rom. Mem. viii. p. 343, pi. 52. 
f. 14(1901). 

Brazil, Amazons. 

(3) Commotria invexustella. 

Coinmotria inveiiustella Berg, An. Soc. Arg. xix. p. 278 
(1885); Eag. Rom. Mem. viii. p. 343, pi. 36. f. 24. 
S. Brazil ; Argentina. 

(4) Commotria arrhabdella, sp. n. 

5 . Head and thorax pale red-brown ; abdomen whitish tinged 
with red-brown, dorsally fidvous yellow towards base ; pectus 
and legs whitish tinged with red-brown. Fore wing pale flesh- 
red slightly irrorated with dark scales ; a terminal series of slight 
dark points. 

Peru, R. Pachaya, 1 2 type. Ux2}. 24 mm. 

Sect. II. (Tolima.) Antenna? of male serrate. 

(5) *Commotria roseopennella. 

Volusia roseojjennella Hulst, Trans. Am. Ent. 8oc. xvii. p. 206 
(1890); Rag. Rom. Mem. viii. p. 340, pi. 51. f. 3; Dyar, Cat. 
Lep. N. Am. p. 438. 
■ U.S.A., Florida. 

(6) *Commotria obertuuri. 

Tolima oherthurii Rag. Nouv. Gen. p. 41 (1888); id. Rom. 
Mem. viii. p. 341, pi. 38. f 4. 
Colombia, 

(7) * Commotria opacella. 

Anerastia opacella Hulst, Ent. Am. iii. p. 138 (1887); Rag. 
Rom. Mem. viii. p. 341, pi. 36. f. 23; Dyar, Cat. Lep. N. Am. 
p. 438. 

U.S.A., Texas. 



PYRALID^, SUBFAMILY HYPSOTROPINiE. 109 

(8) tCoMMOTRIA PHYCITELLA. 

Toliina jyhycitella Rag. Nouv. Gen. p. 41 (1888) ; id. Rom. Mem. 
viii. p. 341,pl. 44. f. 17. • 
Gold Coast. 

Sect. III. {Mangala.) Antennaa of male laminate and ciliated. 

(9) tCoMMOTRIA TRIPARTELLA, sp. 11. 

2 . Head and thorax purple-pink ; abdomen white, dorsally 
fulvous yellow towards base ; pectus, legs, and ventral surface of 
abdomen white tinged with purplish pink ; a,nal tuft yellow 
below. Fore wing with white costal fascia narrowing to apex 
leaving the costal edge dark towards base, then pinkish, defined 
below by a dark reddish-brown fascia, the inner half of wing 
rose-pink, whiter towards inner margin. Hind wing white. 
IST. China, Pekin {Hughes), 1 $ type. Eccp. 20 mm. 

(10) fCoMMOTRIA NEURIAS, Sp. 11. 

Head and thorax white suffused with red-brown ; abdomen 
white, dorsally tinged with fulvous j^ellow towards base ; pectus 
and legs white tinged with brown. Fore wing white tinged with 
reddish brown, the veins white defined on each side by slight 
dark brown streaks ; the costal area defined below by a brown 
shade; obliquely placed almost medial dark brown points in 
submedian fold and on vein 1 ; a dark point just above lower 
angle of cell ; obliquely placed postmedial dark points on veins 6 
to 2 and in submedian fold and a point nearer the teimen 
on vein 1 : a terminal series of black points. Hind wing white 
slightly tinged with ochreous brown. 

N. Nigeria, Zungeru [Macfie), 1 c^ : Br. C. Africa, Mt. Mlanje 
[Neave), 1 J., 1 $ type. Exp. 16 mm. . 

(11) tC0MM0TR,IA ERYTHKOGRAPTA, Sp. n. 

:f . Head and thorax white tinged with rufous; abdomen 
white tinged with ochreous j'ellow ; pectus and legs white mixed 
with red-brown. Fore wing white tinged with rufous, the veins 
and submedian fold white defined on each side by slight rufous 
streaks ; the costal area defined below by a dark brown streak 
and a dark streak below basal half of median nervure ; obliquely 
placed almost medial dark points below the cell and on vein 1 ; a 
dark point just above lower angle of cell and obliqufely placed 
postmedial dark points on veins 6 to 2. Hind wing white with 
a slight ochreous tinge. 

Br. C. Africa, Katungas {de Jersey), 1 S type. Exp. 20 mm. 

(12) tCOMMOTRIA RUFIMEDIA, Sp. U. 

S . Head and thorax white tinged with red-brown ; abdomen 
creamy white. Fore wing white faintly tinged with I'ufous and 
irrorated with fuscous ; a rufous shade below median nervure ; 



1 10 SIH GEORGE HAMPSOX ON THE 

a minute antemedial black spot on vein 1 and minute discoidal 
spot; an oblique postmedial series of black points on veins 4 to 2 
and a point on vein 1 ; a terminal series of black points. Hind 
wing white tinged with ochreous. 

Bb,. C. Africa, Mt. Mlanje (Neave), 1 6 type. Exp. 16 mm. 

(13) tCoMMOTRIA RHODONEURA, Sp. n. 

$. Head and thorax rufous; abdomen whitish tinged with 
rufovis ; palpi whitish below ; fore legs red-brown ; mid and hind 
legs whitish tinged with rufous. Fore wing yellowish white, the 
veins streaked with purplish pink and the costal area sufiused 
with purplish pink; a black antemedial point on vein 1, a 
discoidal point and postmedial points on veins 4, 2, 1 ; a terminal 
sei'ies of black points. Hind wing white tinged with ochreous 
rufous. 

Transvaal, White R. {Cooke), 1 5 type. Exp. 20 mm. 

(14) tCoMMOTRIA rufidelineata, sp. n. 

Head and thorax whitish suffused with iTifous ; abdomen 
creamy wdiite, dorsally fulvous yellow towaixls base ; pectus and 
legs white tinged with red-brown. Fore wing ochi-eous white, 
the veins defined on each side by fine rufous streaks, the costal 
area defined below by a slight rufous shade ; two antemedial 
blackish points on vein 1 and obliquely placed postmedial points 
on veins 4 to 1 ; a terminal series of black points. Hind wing 
ochreous white. 

Br. E. Africa, Nairobi {Anderson), 1 $ type; Br. C. Africa, 
Mt. Mlanje {Neave), 1 S ■ Exp. 24 mm. 

(15) tOOMMOTRIA MIOSTICTA, Sp. n. 

c5' . Head and thox'ax white tinged with rnfous ; abdomen 
creamy white, dorsally fulvous 3'ellow towards base ; pectus, legs, 
and ventral surface of abdomen white suffused with brown. Fore 
wing white tinged witli rufous, the veins white defined on each 
side by fine rufous streaks ; the costal edge blackish towards base ; 
the costal area, defined below by a slight red-bi'own shade ; a 
black point just above lower angle of cell. Hind wing white 
slightly tinged with ochreous. 

Sierra Leone {Clements), 1 S type. Exp. 22 mm. 

(16) *Commotria crassiscapella. 

Mangala crassiscapella Rag. ISTouv. Gen. p. 41 (1888) ; id. Rom. 
Mem. viii. p. 342, pi. 37. f. 17. 
Sudan. 

(17) tCoMMOTRIA phcenicias, sp. n. 

J . Head and thorax whitish suffused with rufous ; abdomen 
yellowish white ; pectus and legs whitish suffused with brown. 
Fore wing white very thickly irroi^ated with deep purple-pink, 



PYRALID^, SUBFAMILY HYPSOTROPIN^. Ill 

the lower part of cell and the area just beyond it much whiter, 
the costa tinged with brown ; a small black-brown discoidal spot ; 
the veins towards termen with slight brown streaks. Hind wing 
yellowish white, the costa tinged with brown towards apex. 

N. Nigeria, Zungeru {Macjfie), 1 d" ; Uganda, Katesa (Betton), 
1 (S type. Exp. 20-22 mm. 

(18) tCoMMOTRIA ROSELLA, Sp. n. 

Head and thorax bright rose-pink ; abdomen ochreous ; pectus 
whitish ; legs and abdomen pink. Fore wing bright rose-pink 
mixed with some whitish except on costal area, the median 
nervure and veins beyond the cell with tine deep pink streaks ; a 
small deep pink spot oi\ upper discocellular ; cilia fuscous at apex. 
Hind wing pale ochreous, the costa and cilia at apex tinged with 
pink. 

Ab. 1. Fore wing with a dark reddish-brown shade along sub- 
costal nei'vure and thence to apex, the spot on upper discocellular 
dark brown. 

Br. C. Africa, Mt. Mla.nje (A'^eave), 4 J , 5 $ type. Exj). 
22-26 mm. 

(19) tCOMMOTRIA ALBINERVELLA, Sp. n, 

2 i Head and thorax pale purplish pink ; abdomen ochreous 
white, dorsally fulvous yellow towards base ; pectus, legs, and 
ventral surface of abdomen whitish tinged with pink. Fore wing 
pale purple-pink, the veins streaked with white ; the costal area 
white tinged with pink, narrowing to apex and defined below by 
a slight brown shade ; a slight dark point at upper angle of cell. 
Hind wing white with a slight ochreous tinge. 

Rhodesia, Bulawayo {Eyles), 1 5 type. Exp. 20 mm. 

(20) tCoMMOTRIA VENOSELLA, Sp. n. 

S • Head, thorax, and abdomen whitish suffused with rufous : 
fore femora and tibife black-brown in front. Fore wing whitish 
suffused with purplish rufous and sliglitly irrorated with blackish 
in the interspaces ; the veins prominently streaked with white. 
Hind wing ochreous whitish, with a slight reddish-brown terminal 
line except towards tornus. 

Br. C. Africa, Mt. Mlanje {JSfeave), 1 S type. Exj). 30 mm. 

(21) fCOMMOTRIA RHODGCHROA, Sp. n. 

S . Head and thorax pale I'ose-pink ; abdomen ochreous white, 
dorsally fulvous yellow towards base ; pectus, legs, and ventral 
surface of abdomen whitish suffused with red-brown. Fore wing 
pale rose-pink, the costal area with a slight red-brown tinge ; the 
veins streaked with white and defined on each side by fine deeper 
pink streaks. Hind wing ochreous white. 

Natal, Tugela R., Bonds' Diift [Reynolds), 1 6 type. Exp. 
26 nim. 



112 SIR GBORGE IIAMPSON ON THE 

(22) tCoMMOTRIA CASTANEIPARS, Sp. n. 

5 . Head, thorax, and abdomen deep red-brown ; hind tibiae 
white towards base. Fore wing with narrow white costal fascia 
narrowing to a point before apex, the area below it deep chestnnt- 
brown to median nervure and vein 2, the inner area purplish 
pink. Hind wing i-ed- brown. 

Br. C. Africa, Mt. Mlanje (A'^eave), 1 $ type. Uxp. 22 mm. 

(23) tCOMMOTRIA PROPH^ELLA, Sp. n. 

5 . Head and thorax pale flesh-red ; abdomen ochreous ; palpi 
tinged with brown ; pectus and legs whitish suffused with brown. 
Fore wing pale flesh-red, the costal area broadly suffused with 
brown, extending on basal half to median nervure ; the veins 
beyond the cell finely streaked with brown. Hind wing ochreous 
white, 

Br. 0. Africa, Mt. Mlanje (JVeave), 1 $ type. " Exp. 22 mm. 

(24) tCOMMOTRIA PHLEBICELLA, Sp. n. 

5 . Head and thorax pale flesh-pink mixed with some whitish ; 
abdomen whitish, dorsally tinged Avith fulvous-yellow towai-ds 
base ; pectus, legs, a,nd ventral surface of abdomen white tinged 
with brown. Fore wing pale flesli-pink, the costal area broadly 
suffused with brown, extending to the median nei-vure towards 
base ; the costal edge and veins finely streaked with white. Hind 
wing ochreous white. 

Mashonaland, Salisbuiy {Marshall), 2 9 type. Exp. 26 mm. 

(25) tCoMMOTRIA eneryella, sp. n. 

d . Head and thorax whitish suffused with pale flesh-red ; 
abdomen ochreous white. Fore wing whitish suffused with pale 
purplish pink and the costal half tingerl with brown ; the veins, 
except on inner area,, stieaked with white and. defined on each 
side by fine brown sti'eaks. Hind wing white tinged with bi^own 
except towards base and inner n^rgin 

Formosa, Takow (Wileman), 1 (S type. Exp. 18 mm. 

Genus SiBOGA. 

Type. 
Sihoga Hmpsn. Rom. Mem. viii. p. 338 (1901) falsella. 

Proboscis aborted and minute ; palpi upturned, the 2nd joint 
reaching to vertex of head and hollowed out to receive the brush- 
like maxillary palpi, the 3i-d joint moderate and porrect ; frons 
with conical prominence ; antennas of male typically serrate and 
ciliated, the basal joint elongate, the shaft with double riflge of 
scales at base above enclosing a hollow. Fore wing long and 
narrow, the apex rounded, the term en obliquely curved ; vein 3 
from close to angle of cell; 4, 5 strongly stalked ; 6 froixl below 
upper angle; 8, 9, 10 stalked; 11 from cell. Hind wing with 



PYRALID.E, SUBFAMILY HYPSOTROPIN^. 113 

vein 2 from well before angle of cell ; 3 and 5 stalked, 4 absent ; 
6, 7 shortly stalked ; 8 not anastomosing with 7. 

Sect. I. Antennae of male unipectinate with moderate branches, the apical part 
ciliated. 

(1) SiBOGA ALBIMEDIELLA, Sp. n. 

Head and thorax white tinged with pink and brown ; abdomen 
ochreous, the first three segments ochreous on dorsum. Fore 
wing pale pink, the veins streaked with white ; the costal ai'ea 
tinged with brown ; a white fascia fi-om base to termen above 
median nervure and vein 5. Hind wing white slightly tinged 
with ochi-eous especially towards termen. 

Punjab, Simla, 1 c5" type. Exp. 26 mm. 

Sect. II. {Siboga.) Antennte of male serrate and ciliated. 

(2) *SlBOGA FALSELLA. 

Hypsotropha falsella Snell. Midden-Sumatra Lep. p. 82 (1880) ; 
Rag, Rom. Mem. viii. p. 338, pi. 38. f. 3. 
Sumatra. 

Sect. III. Antennas of male laminate and almost simple. 

(3) tSlBOGA DIALEUCELLA, Sp. n. 

<S . Head and thorax white, suffused with rufous except on 
vertex of head and dorsum of thorax ; abdomen creamy white, 
dorsally fulvovis yellow towards base ; palpi white in front ; pectus 
and legs white slightly tinged with red-brown. Fore wing white 
suffused with rufous ; the veins white defined on each side bj' 
fine rufous streaks, the median nervure defined b^w by a rufous 
fascia ; a white fascia through the cell, then narrower along 
discal fold to termen. Hind wing ochreous white. 

Kashmir, Goorais Yalley {Leech), 1 S type, Exj). 24 mm, 

(4) tSlBOGA ZEAVORA, Sp. n. ^ 

$ . Head, thorax, and abdomen whitish suffused with rufous, 
the last dorsally fulvous yellow towards base. Fore wing whitish 
tinged with rufous, the veins defined on each side by streaks 
formed of red-brown scales, the cell with two streaks in it ; the 
inner margin irrorated with red-brown ; a terminal series of 
prominent black points. Hind wing ochreous white with a 
terminal series of brown stria? except towards tornus, 

Malay States, 1 $ type. Exp. 30 mm. 

Larva feeds on maize. 



Proc. Zool. Soc— 1918, No. YIII. 



114 SIR GEORGE HAMPSOt? ON THE 

Genus Ematheudes. 

Type. 

Ematheudes ZeU. Stett. Ent. Zeit. 1867, p. 385 ... panctella. 

Proboscis aborted and minute ; palpi downcurved, extending 
about three times length of head and thickly scaled ; maxillary 
palpi minute and filiform ; frons with large tuft of scales ; 
antennse of male minutely sei'rate and ciliated. Fore wing 
narrow, the apex rounded, the termen evenly curved ; vein 3 
from close to angle of cell ; 4, 5 stalked ; 6 from below upper 
angle; 8, 9, 10 stalked; 11 from cell. Hind wing with vein 2 
fi*om well before angle of cell ; 3 and 5 stalked, 4 absent ; 6, 7 
shortly stalked ; 8 not anastomosing with 7. 

(1) Ematheudes punctella. 

Chilo punctella Treit. Schmett. Eur. ix. 2, p. 268 (1833) ; Dup. 
Lep. Fr. pi. 273. f. 4 ; Herr.-Schiiff. Eur. Sclimett. iv. p. 108, 
Tin. f. 85 ; Rag. Rom. Mem. viii. p. 333 ; Htaud. Oat. Lep. pal. 
ii. p. 13. 

tS. France ; Spain ; Corsica ; Italy ; Sicily ; Morocco ; 
Greece ; Turkey ; Cyprus ; Asia Minor ; Syria. 

(2) *Ematheudes pseudopunctella. 

Ematheudes jDseudojnmctella Rag. Nouv. Gen. p. 40 (1888) ; id. 
Rom. M6m. viii. p. 334, pi. 36. f. 22 ; Stand. Cat. Lep. pal. ii. p. 13. 
Syria. Probably an aberration of E. punctella'. 

(3) Ematheudes straminella. 

Ematheudes straminella Snell. Tijd. v. Ent. 1872, p. 107, pi. 8. 
f. 9. 

Gambia ; Sierra Leone ; Angola ; Br. C. Africa ; Portu- 
guese E. Africa. 

(4) fEMATHEUDES LENTISTRIGALIS. 

Emmatheudes lentistrigalis Hmpsn. Tr. Zool. Soc. xix. p. 134, 
pi. iv. f. 65 (1909). 

Gold Coast ; N. Nigeria ; Br. E. Africa ; Uganda ; Br. C. 
Africa. 

(5) Ematheudes paleatella. 

Einatheudes puleateila Rag. Nouv. Gen. p. 40 (1888); id. Rom. 
M6m. viii. p. 334, pi. 36. f. 20. 

Br. E. Africa; Uganda; Br. C. Africa; Mashonaland ; 
Transvaal ; Natal ; Basutoland ; Cape Colony. 

(6) Ematheudes tunesiella. 

Ennatheudes tunesiella Rag. Iris, v. p. 298 (1892); id. Rom. 
Mem. viii. p. 335, pi. 42. f. 22; Stand. Cat. Lep. pal. ii. p. -13. 
S, Italy ; Tunis ; Syria ; W. Turkestan, 



PYRALID.E, SUBKAMILY HYPSOTROPINiE. 115 

\ 

(7) Ematheudes crassinotella. 

Ematheudes crasshiolella Rag. Nouv. Gen. p. 41 (1888); id. 
Rom. Mem. viii. p. 335, pi. 35. f. 26. 
Zanzibar ; Br. E. Africa ; Mashonaland ; Natal. 

(8) *Ematheudbs varicella. 

Ematheudes varicella Rag. Ann. Soc. Ent. Fr. 1887, p. 258 ; 
id. Rom. Mem. viii. p. 336, pi. 35. f. 27; Stand. Oat. Lep. pal. 
ii. p. 13. 

Armenia ; W, Turkestan. 

(9) *Ematheudks vitellinella. 

Etnathevdes vitellinella Rag. Ann. Soc. Ent. Fr. 1887, p. 258; 
id. Rom. Mem. viii. p. 336, pi. 49. L 22; Stand. Cat. Lep. pal. 
ii. p. 13. 

Asia Minor, Georgia. 

(10) *Ematheudes euchlytella. 

Ematheudes euchlytella Rag. Nonv. Gen. p. 41 (1888) ; id. 
Rom. Mem. viii. p. 337, pi. 36. f. 21. 
Argentina. 

Genus Biafra. 

Type. 
Biafra Rag. Nouv. Gen. p. 40 (1888) concinnella. 

Proboscis aborted and minute ; palpi downcurved, extending 
about three times length of head and moderately scaled ; maxil- 
lary palpi slightly dilated with scales ; irons smooth, with large 
pointed tuft of hair; antenna? of male minutely serrate and 
ciliated, the basal joint rather long, the shaft with double ridge 
of scales at base enclosing a hollow. Foi-e wing very narrow, 
the apex rounded, the termen obliquely curved ; vein 3 from near 
angle of cell ; 4, 5 separate ; 6 from below upper angle; 8, 9, 10 
stalked ; 11 from cell. Hind wing with vein 2 from near angle 
of cell ; 3 and 5 stalked, 4 absent ; 6, 7 shortly stalked ; 8 anas- 
tomosing with 7. 

(1) Biafra concinnella. 

Biafra concinnella Rag. Nouv. Gen. p. 40 (1888); id. Rom. 
Mem. viii. p. 330, pi. 38. f. 2. 

N. Nigeria ; Br. C. Africa ; Mashonaland ; Transvaal ; 
Natal. 

(2) tBiAFRA RHODINELLA. 

Biafra rhodinella Rag. Nouv. Gen. p. .40 (1888); id. Roin. 
Mem. viii. p. 381, pi. 44. f. 20. 

Gold Coast; N.Nigeria; Mashonaland; Transvaal. 



116 SIR GEORGE HAMPSON ON THE 



Genus Ethiotropa, nov. 

Type, E. 'pyromerella. 

Pi-oboscis aborted and minute ; palpi porrect and slightly 
downcurved, extending about twice the length of head, the 2nd 
joint fringed with rough scales below, the 3rd moderate ; maxil- 
lary palpi strongly dilated with scales; frons smooth, with 
pointed tuft of hair above ; antennte of male somewhat laminate 
and minutely cihated, the basal joint long, the shaft with ridge 
of scales above at base. Fore wing long and very narrow, the 
apex rounded, the termen obliquely curved ; veins 3, 4 stalked 
on one side of the specimen from a point on the other ; 5 sepa- 
rate ; 6 from below upper angle; 8, 10, 11 stalked, 9 absent. 
Hind wing with vein 2 from Avell before angle of cell ; 3 and 5 
from angle and approximated for some distance, 4 absent ; 6, 7 
shortly stalked ; 8 anastomosing with 7. 

Ethiotropa pyromerella, sp. n. 

S . Head, tegulae, and base of patagia fiery red, the rest 
of thorax brownish ochreous ; abdomen ochreoas white, dorsally 
fulvous yellow towards base ; pectus, legs, and ventral surface of 
abdomen whitish suffused with brown. Fore wing with whitish 
costal fascia narrowing to apex, the costal edge fiery red to 
beyond middle, defined below by a black-brown fascia ; the rest 
of wing fiery red. Hind wing v/hite tinged with ochreous 
brown. 

N. Nigeria, Akassa (^Lugard), 1 c? type. Exp. 22 mm. 



Genus Baptotropa, nov. 

Type, B. tricolorella. 

Proboscis aborted and minute ; palpi porrect and slightly 
downcurved, extending about three times length of head, the 
2nd joint fringed with rough scales above, the 3rd moderate ; 
maxillary palpi minute and filiform ; frons smooth and rounded, 
without tuft of hair ; antennfe of male strongly uniserrate, the 
basal joint long, the shaft with double ridge of scales at base 
enclosing a hollow. Fore wing narrow, the apex rounded, the 
termen evenly curved ; vein 3 from before angle of cell ; 4, 5 
from angle; 6 from below upper angle; 8, 9, 10 stalked; 11 
from cell. Hind wing with vein 2 from well before angle of 
cell ; 3 and 5 from angle, 4 absent ; 6, 7 shortly stalked ; 8 
anastomosing with 7. 

fBAPTOTROPA TRICOLORELLA. 

Patna tricolorella Hmpsn. J. Bomb. N. H. Soc. xii, p. 308 
(1899); Rag. Rom. Mem. viii. p. 340, pi. 52. f. 15. 
Assam, Khasis. 



PYRALID^, SUBFAMILY HYPSOTROPIN^. 117 

Genus Patna. 

Type. 

Patna Rag. Nouv. Gen. p. 39 (1888) ehoricostella. 

Proboscis absent ; palpi poiTecfc and almost straight, extending 
about three times length of head, the 2nd joint slightly fringed 
with hair above towards extremity, the 3rd moderate ; maxillary 
palpi small and filiform ; irons smooth and rounded, without 
tuft of hair ; antennae of female almost simple. Fore wing 
rather narrow, the costa tyj)ically almost straight, the apex 
rounded, the termen evenly curved ; vein 3 from near angle of 
cell ; 4, 5 from angle and aj)proximated for a short distance ; 6 
from below upper angle; 8, 9, 10 stalked ; 11 from cell. Hind 
wing with vein 2 from well before angle of cell ; 3 and 5 from 
angle, 4 absent ; 6, 7 from upper angle ; 8 not anastomosing 
with 7. 

(1) Patna ehoricostella. 

Patna ehoricostella Rag. Nouv. Gen. p. 39 (1888); Hmpsn. 
Moths Ind. iv. p. 57 ; Rag. Rom. Mem. viii. p. 330, pi. 38. f. 1. 
SiKHiM ; Bhutan. 

(2) t Patna venatella, sp. n, 

5 . Head and thoiax pale rufous ; palpi whitish except above ; 
pectus, legs, and abdomen whitish tinged with red-brown. Fore 
wing white, the veins, discal fold in the cell and the submedian 
fold defined by fine purplish-pink streaks ; a minute dark brown 
spot just above lower angle of cell ; a terminal series of black 
points. Hind wing white slightly tinged with ochreous. 

Br. E. Africa, Tagas (Betton), I $ , Aios {Betton), 1 $ type. 
Exp. 32 mm. 

(3) fPATNA BRUNNEICOSTELLA, Sp. n. 

2 . Head and thoi'ax pale flesh-pink ;. abdomen ochreous 
white ; palpi, pectus, legs, and ventral surface of abdomen 
whitish suffused with red-brown. Fore wing white, the costal 
area tinged with brown, the area below the cell and vein 2 
sufli"used with pale pink ; the veins white, those beyond the cell 
defined on each side by fine pink streaks, the inedian nervure and 
vein 2 defined below by stronger sti'eaks ; a dark point in lower 
angle of cell. Hind wing ochreous white. 

Transvaal, White H. {Cooke), 1 $ type. Exp. 32 mm. 

Genus MEaALOPHOTA, nov. 

Type, M. leonella. 

Proboscis absent ; palpi obliquely upturned, the 2nd joint 
reaching. to above vertex of head, dilated and hollowed out to 
receive the brush-like maxillaiy palpi, the 3rd minute ; frons 



118 SIR GEORGE HAMPSON ON THE 

with long truncate conical prominence ; antennae of male pecti- 
nate with rather long uniseriate branches to near apex, the shaft 
with large sinus at base containing a double ridge of scales 
enclosing a hollow. Fore wing rather long and narrow, the apex 
rounded, the tei-men evenly curved ; vein 2 from well before 
angle of cell ; 3 from before angle ; 4, 5 from angle ; 6 from 
below upper angle; 8, 9 stalked; 10, 11 from cell.' Hind wing 
with vein 2 from well before angle of cell ; 3 and 5 stalked, 
4 absent ; 6, 7 shortly stalked ; 8 not anastomosing with 7. 

MeGALOPHOTA LEON ELLA, Sp. n. 

d . Head and thorax ochreous white with a slight red-brown 
tinge on shoulders, the antennal tufts black on inner side; 
abdomen ochi-eous ; pectus and legs ochreous white tinged with 
red-brown. Fore wing ochreous white irrorated with brown, 
the costal area slightly irrorated to neai' apex. Hind wing 
ochreous white. 

Sierra Leoxe [Dadyeon), 1 cJ type. Uxp. 20 mm. 

Genus Martia. 

Type. 

Martia Rag. N. Am. Phyc. p. 18 (1887) arizonella. 

Ur'ula Hulst, Can. Ent. xxxii. p. 175 (1900) arizonella. 

Proboscis absent ; palpi with the 2nd joint porrect, extending 
about twice the length of head, the 3rd I'ather oblique, long, 
slender, and somewhat acute at extremity ; mnxillaiy palpi 
slight and filiform ; froiis with large truncate conical prominence; 
antennjfi of male minutely serrate and with fascicles of long cilia. 
Fore wing rather narrow, the apex rounded, the termen evenly 
curved ; vein 2 from long before angle of cell ; 3 from before 
angle; 5 from above angle; 6 from below vipper angle; 8, 9 
stalked; 10, 11 from cell. Hind wing with vein 2 from long 
before angle of cell ; 3 and 5 from angle, approximated for a 
short distance, 4 absent ; 6, 7 from upper angle ; 8 not anasto- 
mosing with 7. 

Martia arizonella. 

Martia arizonella Rag. N. Am. Phyc. p. 18 (1887) ; id. Rom. 
Mem. viii. p. 367, pi. 38. f. 20 ; Dyar, Cat. Lep. N. Am. p. 489, 

Urida incongruella Hulst, Can. Ent. xxxii. p. 175 (1900); 
Dyar, Cat. Lep. N. Am. p. 437. 

U.S.A., Colorado, Arizona. 

Genus DiSCOFRONTIA, 

Type. 
DisGofrontia Hm})sn. Rom. Mem. viii. p. 350(1901). normella. 

Pioboscis aborted and minute ; palpi upturned, the 2nd joint 
reaching to vertex of head and moderately scaled and flattened. 



PYRALID.E, SUBFAJIILY HYPSOTROPIN.E. 110 

the 3i'd moderate; maxillary palpi slight and filiform; frons 
broad, with a disk of concentric white scales converging to middle; 
antennfe of male strongly serrate, with a large sinus and ridge of 
scales at base of shaft. Fore wing rather narrow, the apex 
rounded, the termen evenly curved;. vein 2 from Avell before 
angle of cell ; 3 fi-om before angle ; 4, 5 from angle : 6 from 
below upper angle; 8, 9 stalked; 10, 11 from cell. Hind wing 
with vein 2 fi-om before angle of ce\l ; 3 and 5 from angle, 
4 absent ; 6, 7 from upper angle ; 8 not anastomosing with 7. 

*DlSCOFRONTlA NOKMELLA. 

Discofrontia normella Hmpsn. Rom. Mem. viii. p. 350, pi. 52. 
f. 20 (1901). 
Natal. 

Genus Critonia. 

' . . . . Type. 

Critoma Rag. Bull. Soc. Ent. Fr. 1890, p. ccxiv. suhconcinnella. 

Singhalia Hmpsn. J. Bomb. Nat. Hist. Soc. xii. 
p. 309 (1898) ; id. Rom. Mem. viii. p. 351 
(1 901 ) sarcoylauca. 

Proboscis aborted and minute ; palpi typically downcurved, 
extending about three times length of head and moderately 
scaled ; maxillaiy palpi small and filiform ; frons. smooth and 
with slight tuft of hair ; antennse of male typically serrate and 
fasciculate, the shaft with sinus fvt base containing a large ridge 
of scales. Fore wing rather long and narrow, the apex rounded, 
the termen evenly curved ; vein 3 from before angle of cell ; 
4, 5 from just above angle ; 6 from below upper angle ; 8, 9 
stalked ; 10, 1 1 from cell, 10 approximated to 8, 9. Hind wing 
with veins 3 and 5 fi'om angle of cell and approximated for a 
short distance, 4 absent ; 6, 7 shortly stalked ; 8 not anastomos- 
ing with 7. 

Sect. I. Palpi of male obliquely upturned, the 2nd joint hollowed out to receive 
the brush-like maxillary palpi, the 3rd short; antennsB laminate, with large 
sinus and ridge of scales at base of shaft. 

(1) fCRITONIA PH^ONEURA, Sp. n. 

(S . Head and thorax whitish sufiused with I'ed-biown ; abdo- 
men ochreous white, dorsally fulvous yellow towards base ; 
pectus and legs whitish tinged with brown. .l*'ore wing whitish 
tinged Avith red-brown, the veins except on inner area blackish 
defined on each side by fine white streaks ; the costal area white, 
narrowing to apex and defined below by a red-brown shade, the 
costal. edge brown to beyond middle and the veins on costal area 
finely streaked with black ; a, minute a.ntemedial black spot on 
vein l,a point at middle of submedian fold and minute post- 
medial streaks above vein 2 and on vein 1 ; cilia flesh-white with 



120 Sir GEORGE hampson on the 

series of minute black streaks near base except towards tornus. 
Hind wing whitish tinged with brown especially on costal area 
and at termen, 

Formosa, Banshorio {Wile7nan), 1 S type. Exjj. 18 mm. 

(2) tORITONIA PROMEL^NA. 

Critonia promelo&na Hmpsn. J. Bomb. Nat. Hist. Soc. xii. 
p. 309 (1898) ; id. Rom. M6m. viii. p. 366, pi. 51. f. 23. 

SiKHIM. 

(3) tORITONIA PURPUREOTINCTA. "" 

Critonia pibrpureotincta Hmpsn. Moths Ind. iv. p. 61 (1896); 
id. Rom. Mem. viii. p. 365, pi. 51. f. 22. 
SiKHiM; Bhutan. 

(4) tCRITONIA HOLORHODA. 

Critonia holorhoda Hmpsn. J. Bomb, Nat. Hist. Soc. xviii. 
p. 259(1908). 
Ceylon. 

Sect. II. Maxillary palpi of male filiform. 

A. [Critonia). Aiiteniue of male serrate, with larger sinus and ridge of scales 
at base of shaft ; palpi downcurved and about three times length of head. 

(5) *Critonia sltbconcinnella. 

Critonia subconcinnella'Rfig. Bull. Soc. Ent. Fr. 1890, p. ccxiv : 
id. Rom. Mem. viii. p. 365, pi. 6. f. 20. 
Burma. 

(6) fORITONTA LEUCOPLEURA, Sp. n. 

J . Head and thorax whitish suffused with purplish pink ; 
abdomen whitish suffused with ochreous brown ; antennte with 
the tuft blackish ; palpi, pectus, and legs whitish tinged with 
brown. Fore wing whitish suffused with purplish pink, the 
veins streaked with blackish, vein 1 only towards termen ; the 
costal area pure white, narrowing to apex and defined below by 
a blackish shade ; the costal edge dark to beyond middle and the 
interspaces of terminal area Avith slight dark streaks except 
towards tornus. Hind wing whitish, the costal area broadly 
suffused with bi'own. 

Basutoland, Maseru (Crawshai/), 1 J type. Exp. 26 mm. 

(7) tCRITONIA ROSEISTRIGELLA. 

Critonia roseislrigella Hmpsn. Moths Ind. iv. p. 61 (1896) ; id. 
Rom. Mem. viii. p. 365, pi. 51. f. 21. 

Madras, Nilgiris ; Philippines, Luzon. 



PYRALID^, SUBFAMILY HYPSOTROPIN^. 121 

(8) Critonia OCHRACEALIS. 

Critonia ochracealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi. 
p. 1251 (1912). 

Punjab, Kangia ; Madras, Nilgiris. 

(9) *Oritonia hilgerti. 

Pectinigeria hilgerti Roths. Nov. Zool. xxii. p. 236. 
Algeria. 

B. Antennas of male laminate and without sinus and ridge of scales at base of 
shaft. 

a. (Sinffhalia.) Palpi of male with the 2nd joint obliquely upturned to vertex 

of head and thickly scaled, the 3rd porrect, long and blunt. 

(10) tORITONIA SARCOGLAUCA. 

Critonia sarcoglauca Hmpsn. Moths Inch iv. p. 60 (1896); id. 
Rom. Mem. viii. p. 351, pi. 51. f. 24. 
Ceylon. 

b. Palpi of male downcurved and extending about three times length of head. 

(11) Critonia rhodessa. 

Saluria rhodessa Turner, Pr. R. Soc. Queensh xviii. p. 120 
(1903). 

Queensland. 

(12) tCRITONIA SARCOIDA, Sp. n. 

Head n.nd thorax purplish pink, the head and shouldeis 
tinged with brown ; abdomen ochreous white, dorsally fulvous 
yellow towards base ; pectus, legs, and ventral surface of abdo- 
men ochreous tinged with brown. Fore wing purpli&li pink, the 
costal area tinged with biown ;" the veins, discal fold iii the cell, 
and submedian fold white defined on each side by fine puiplish- 
pink streaks, the median nervure rather more stiongly stieaked 
with white and with a brown streak below it. Hind wing 
ochreous white, the costal area tinged with brown. 

Br. E. Africa, Njora {Chohnley), 1 S ; Br. C. Africa, 
Mt. Mlanje [Neave), 3 $ ; Portuguese £. Africa, Kola Valley 
(iVertve), 3 § ; Mashonaland, Salisbury (i)/«rsAaZ^), 1 $ ; Trans- 
vaal {Fead), 1 J type. Hxjj. 28-34 mm. 

Genus MONOCTENOCERA. 

Type. 
Monoctenocera Hmpsn. J. Bomb. Nat. Hist. iSoc. xii. 

p. 310 (1898) ; id. Rom. Mem. viii. p. 311 (1901). hrachiella. 

Proboscis aborted 8nd minute ; palpi obliquely upturned, the 
2nd joint reaching to just above vertex of head and hollowed out 
to receive the brush-like maxillary palpi, the 3i'd short and 



122 SIR GEORGE IIAMPSON ON THE 

thickly scaled ; frons suiooth, with large tuft of hair ; antennte 
of male unipectiuate, typically wibli very short branches, the 
apical part serrate, the shafh with large siaus and ridge of scales 
at base ; mid and hitid tibiiB typically fringed with long hair. 
Fore wing narrow, the apex rounded, the termen evenly curved ; 
vein 3 from close to angle of cell ; 4, 5 strongly stalked ; 6 
from below upper angle; 8, 9, 10 stalked; 11 from cell. Hind 
wing with vein 3 closely approximated to 4, 5 for some distance ; 
4, 5 strongly stalked ; 6, 7 shortly stalked ; 8 anastomosing 
with 7. 

Sect. I. Aiiteniiffi of male with tlie brandies long- ; mid tibiae fringed with hair at 
base only, the hind tibial at extremity only. 

(1) MONOGTEXOCERA LEUCANIA. 

Catagela leucania Feld. Reis. ISTov. pi. 137. f. 13 (1874); 
Hmpsn. Moths Ind. iv. p. 63 ; id. Rom. Mem. viii. p. 312, pi. 51. 
f. 17. 

Hah. Madras, Nilgiris ; Travancore ; Ceylox. 

Sect. II. Antenna; of male with short branches towards base, then serrate : mid and 
hind tibiae fringed with long hair throughout. 

(2) Monoctenocera BRAOHIELLA. 

Polyocha brachiella Hmpsn. J. Bomb. Nat. Hist. 8oc. xii. 
p. 310 (1898) ; id. Rom. Mem. viii. p. 312, pi. 36. f, 6. 
Hah. SiKKiM ; Bencal, Calcutta; Borneo. 



Genus Saborma. 

Type. 
Sahorma Rag. Nouv. Gen. p. 37 (1887) forcipella. 

Proboscis aborted, minute ; palpi of male upturned to about 
vertex of head, slender, typically hollowed out to contain the 
brush-like maxillary palpi ; frons smooth ; antennte of male 
typically strongly serrate, with sinus and large double ridge of 
scales at base. Fore wing narrow, the apex rounded, the termeu 
evenly curved ; vein 3 from close to angle of cell ; 4, 5 sti-ongly 
stalked; 6 from below upper angle; 8, 9, 10 stalked; 11 from 
cell. Hind wing with vein 3 approximated for .some distance to 
4, 5 which are strongly stalked ; 6, 7 shortly stalked; 8 not 
anastomosing with 7. 

Sect. I. Palpi of male with the 2nd joint hollowed out to receive the brush-like 
maxillary palpi ; antennas serrate. 

(1) *8aborma forcipella. 

Sahorma forcipella Rag. Nouv. Gen. p. 37 (1888); id. Rom. 
Mem. viii. p. 310, pL 35. f. 22. 
Sumatra. 



PyUALlD.E, SUBFAStlLY HY PSOTROPINJi. 123 

(2) *Saborma vicina. 

Anerastia vicina Saalm. Ber. Senck, Ges. 1879, p. 307 ; id. Lep. 
Madag. p. 511 ; Rag. Rom. Mem. viii. p. 309, pi. 42. f. 21. 
Madagascar. 

Sect. II. Maxillary palpi of male filiform; antemiie laminate and ciliated. 

(3) fSABORMA PAPUACOLA, Sp. n. 

S . Head and thoi'ax whitish tinged with led-browii ; abdomen 
whitish tinged with brown, dorsally fulvous yellow towards base ; 
antennas with the tuft blackish on inner side ; pectus and legs 
whitish tinged with brown. Fore wing white tinged with 
ochreous brown, the veins, discal fold in the cell, and the sub- 
median fold white defined on each side by streaks formed of 
blackish scales ; a terminal series of black points. Hind wing 
ochreous white with tei-minal series of dark points and stride. 

.Dutch N.. Guinea, Mimika R. {Wollaston), 3 c5' type. Exp. 
26-30 mm. 

Genus Osacia. 
Osakia Rag. Rom. Mem. viii. p. 318 (1901) lineolella. 

Proboscis aborted and minute ; palpi obliquely upturned to 
about vertex of head, moderately scaled; maxillarj^ palpi each a 
minute brush of scales ; frons smooth and rounded ; antenn?e of 
male ciliated, the shaft with sinus and double ridge of scales at 
b.nse ; tibite fringed with hair. Fore wing rather narrow, the 
apex rounded, the termen evenly curved ; veins 2 and 3 stalked 
from before angle of cell : 4, 5 separate ; 6 from below upper 
angle ; 8, 9 stalked ; 10, 11 from cell. Hind wing with vein 3 
from angle of cell ; 4 and 5 strongly stalked ; 6, 7 strongly stalked ; 
8 not anastomosing with 7. 

*0SACIA LINEOLELLA. 

Osakia lineolella Rag. Rom. Mem. viii. p. 319, pi. 43. f. 21 
(1901). 
Japan. 

Genus Ragonotia. 

Tj'pe. 
Ciris Rag. TST. Am. Phyc. p. 17 (1887), nee Grote, 

Lep. 1863 dotalis. 

Ragonotia Grote, Can. Ent. .xx. p. 75 (1888) dotalis. 

Proboscis aborted and minute ; palpi downturned, about three 
times length of head and rather broadly fi'inged with scales 
below ; maxillary palpi filiform ; frons with small lounded pro- 
minence with corneous plate below it ; antennseof male ciliated, 
the basal joint large. Fore wing long and nari'ow, the apex 
rounded, the termen obliquely curved ; vein 2 from towards 
angle of cell ; 3 and 5 from close to angle ; 6 fi-om below upper 



124 SIR GEORGE HAMPSON ON THE 

angle; 8, 9 stalked; 10, 11 from cell. Hind wing with vein 2 
from close to angle of cell ; 3 approximated for some distance to 
4, 5 which are strongly stalked, or 4 rarely absent ; 6, 7 stalked; 
8 not anastomosing with 7. 

Ragonotia dotalis. 

Anerastia dotalis Hiilst, Trans. Am. Ent. Soc. xiii. p. 164 ( 1886) ; 
Rag. Rom. Mem. viii. p. 329, pi. 38. f. 19; Dyar, Cat?. Lep. N. 
Am. p. 437. 

Ciris discigerella Rag. N. Am. Phyc. p. 17 (1887). 

U.S.A., Colorado, Arizona. 

Genus PoLYOCHA. 

Type. 

Folyocha Zeller, Isis, 1848, p. 876 sanguinariella. 

Polyochodes Chretien, Bull. Soc. Ent. Fr. 1911, 

p. 13 stij)ella. 

Proboscis aborted and minute ; palpi typically dovvncurved, 
extending about three times length , of head, the 2nd joint 
moderately scaled, the 3rd rather long and naked ; maxillary 
palpi dilated with scales ; frons smooth, with tuft of scales ; 
antennae of male typically laminate and without sinus and ridge 
of scales at base. Fore wing long and narrow, the apex rounded, 
the termen obliquely curved ; vein 3 from close to angle of cell ; 
4, 5 strongly stalked ; 6 from below upper angle ; 8, 9 stalked ; 
10, 11 from cell. Hind wing with vein 2 from close to angle of 
cell ; 3 approximated for some distance to 4, 5 which are strongly 
stalked ; 6, 7 shortly stalked ; 8 not anastomosing with 7. 

Sect. I. Palpi of male obliquely upturned ; auteiniaa serrate, with sinus and ridge of 
scales at base of shaft. '' 

(1) tPoLYOCHA PLINTHOCHROA, Sp. n. 

Head and thorax bright rufous mixed with some ochreous ; 
abdomen ochreous, dorsally fulvous yellow towards base; antenna? 
ochreous. Fore wing bright rufous with a faint purplish gloss, 
some ochreous in lower part of cell and below and just beyond 
the cell, the veins remaining rufous ; a narrow yellowish- white 
costal fascia, tapeiing to a point just befoi-e apex. Hind wing 
ochreous white, the costal area and termen tinged with red- 
brown. 

Transvaal, Karina (Cooke), I J type. White R, (Cooke), 1 $ . 
Exp. 26 mm. 

(2) tPOLYOCIIA LEUCOPLEURELLA. 

Emmalocera leucopleurella Rag. Nouv. Gen. p. 38 (1888); id. 
Rom. Mem. viii. p. 317, pi. 44. f. 15. 

Gold Coast; S. &. N. Nigeria; Kashmir ; Madras. 



PYRALID^, SUBFAMILY H YPSOTROPIN^E. 125 

(3) tPoLYOCHA GENSANALIS. 

Emmalocera gensanalis South, Trans. Ent. Soc. 1901, p. 405, 
pi. xiv. f. 30. 

COREA. 

Sect. II. Palpi of male downcuvved ; antenna without sinus and ridge of scales at 
base of shaft. 

A. {Folyochodes). Antennae of male pectinate with uniseviate branches. 

(4) *PoLYOCHA STIPELLA. 

Polyochodes stipella Chretien, Bull. Soc. Ent. Fr. 1911, p. 13. 
Algeria. 

B. {Folyoclm). Antenna? of male laminate. 

(5) tPOLYOCHA CINERELLA. 

Polyocha cinerella Hmpsn. Moths Ind. iv. p. 62 (1896) ; id. 
Rom.'^Mem. viii. p. 328, pi. 55. f. 4. 
Punjab ; Bengal. 

(6) Polyocha venosa. 

Epischnia venosa Zell. Isis, 1847,-p. 31 ; Herr.-Schaft'. Schmett. 
Eur. iv. p. 109 ; Rag. Rom. Mem. viii. p.. 327 ; StaurV. Cat. Lep. 
pal. ii. p. 13. 

Cyprus ; Syria. 

(7) Polyocha sanguinartella. 

Polyocha sanguinariella Zell. Isis, 1848, p. 876 ; Rag. Rom. 
Mem.' viii. p. 327, pi. 8. f. 20. 

Br. 0. Africa ; Mashonaland ; Transvaal ; Madagascar. 

(8) Polyocha vesculella. 

Polyocha vesctdella Rag. Nouv. Gen. p. 39 (1888) ; id. Rom. 
Mem. viii. p. 323, pi. 36. f. 11 ; Hmpsn. Moths Ind. iv. p. 63. 
Madras, Palni Hills ; Travancore. 

(9) *Polyocha flagrantella. 

Polyocha Jlagrantella Rag. Rora. Mem, viii. p. 323, pi. 44. 1". 24 
(1901). 

Madagascar. 

(10) fPoLYOCHA STRTGIVENELLA. 

Polyocha strigivenella Hmpsn. J. Bomb. ISTat. Hist. Soc. xii. 
p. 310 (1898) ; id. Rom. Mem. viii. p. 322, pi. 51. f. 19. 
Burma. 



126 SIR GEORGE HAMPSON OX THE 

(11) *POLYOCHA NEUROPTERELLA. 

Polyocha neioropterella Rag, Ann. Soc. Ent. Fr. 1887, p. 258; 
id. Rom. Mem. viii. p. 322, pi. 35. f. 23 ; Stand. Cat. Lep. pal. ii. 
p. 13. 

W. TirRKESTAN. 

(12) *PoLYOCHA FOUCARTl. 

Polyocha foucarti Rag. Ann. Soc. Ent. Fr. 1887, p. 258 ; id. Rom. 
Mem.Ndii. p. 322, pi. 35. f. 24; Stand. Cat. Lep. pal. ii. p. 13. 
Algeria. 

(13) tPOLYOCHA achromatella, sp. n. 

$ . Head and thorax white tinged with ochreous ; abdomen 
white, dorsally f ulvons yellow towards base ; pectns and legs 
ochreous white. Fore wing pale ochreous, the veins white, less 
distinctly so on costal Area. Hind wing white. 

ISr. S. Wales, Broken Hill {Lower), 3 $ type. Exj). 28 mm. 

(14) * Polyocha betritella. 

Polyocha detritella Rag. Nouv. Gen. p. 39(1888); id. Rom. 
Mem. viii. p. 326, pi. 36. f. 14; Hmpsn. Moths Ind. iv. p. 63. 

Punjab: 

(15) fPOLYOCHA FUSCIOOSTELLA, Sp. n. 

5 . Head and thorax glossy fuscous brown ; abdomen fuscous 
brown, pale red-brown at sides and exti-emity ; pectus and legs 
pale red-brown, the" tarsi fuscous brown. Fore wing pale 
red-brown, the costal area broadly glossy fuscous bi'own and 
the inner basal ai-ea tinged with fuscous bi'own. Hind wing 
ochreous white, the costal area tinged with brown. 

N. Nigeria, Zungeru (Alacfie), 2 $ type. Exp. 24 mm. 

Genus Emmalocera. 

Type. 

Eminaloc'era Rag. Nouv. Gen. p. 38 (1888) lei(,cocincta. 

Lodiana Rag. Nouv. Gen. p. 38 (1888) umhriviUella. 

Papua Rag. Bull. Soc. Ent. Fr. 1889, p. ccxx... latilimhella. 

Proboscis aborted and minute; palpi of male t^^picall}^ obliquely 
upturned to above vertex of head, the 2nd joint hollowed out to 
receive the brush-like maxillary palpi, the 3rd short and porrect; 
frons smooth, obliquely flattened ; antennpe of male typically 
with short uniseriate branches, the basal joint large, the shaft 
with large sinus and ridge of scales at base. Fore wing rather 
long and narrow, the apex rounded, the termen evenly curved ; 
vein 3 from near angle of cell, 5 from just above angle ; 6 from 
below upper angle; 8, 9 stalked; 10, 11 from cell. Hind wing 



PYRALIDiE, SUBFAMILY HYPSOTROPIN^. 127 

with vein 2 fi'om well before angle of cell ; 3 npproxiinated for 
some distance to 4, 5 which are strongly stalked ; 6, 7 shortly 
stalked ; 8 not anastomosing with 7. 

Sect. I. Palpi of male upturned, the '2iid joint hollowed out to receive the brush- 
like maxillary palpi ; anteunie with sinus and ridge of scales at base of shaft. 
A. Antennffi of male with rather long uniseriate branches. 

(1) tEMMALOCERA ORNATELLA. 

Polyocha omatella Hmpsn. J. Bomb. Nat. Hist. Poc. xv. p. 21 
(1903). 
Punjab. 

(2) temmalocera pulverealis. 

Polyocha pulverealis Hmpsn. J. Bomb. Nat. Hi.st. Soc. xv. 
p. 20(1903). 

Assam. 

(3) tEMMALOCERA ENDOPYRELLA, Sp. n. 

(S . Head ochreous tinged with piu'plish pink, the antennae 
ochreous ; thorax bright purplish pink ; abdomen fulvous yellow ; 
pectus and legs ochreous tinged with red-brown. Fore wing 
with narrow pure white costal fascia, the rest of wing bi'ight 
purplish pink, suffused with red-brown to median nervure and 
veins 4. Hind wing ochreous white, the costa slightly tinged 
with brown. 

Assam, KhAsis, 1 c? type. Exp. 26 mm. 

(4) Emmalocera loxgiramella. 

Emmalocera longiramella Hmpsn. Rom. Mem. viii. p. 315, 
pi. 52. f. 16 (1901). 
Queensland. 

(5) *Emmalocera radiatella. 

Emmalocera radiatella Hmpsn. Rom. Mem. viii. p. 315, pi. 52. 
f. 21 (1901). 
Queensland. 

(6) tEMMALOCERA ACTINOLEUCA, Sp. n. 

S . Head and thorax pale purplish pink mixed with white : 
abdomen creamy white, dorsally fulvous yellow toAvards base; 
antennae white ; frons, palpi, pectus, and legs white sufliised with 
rufous. Fore wing pale purplish pink, the veins, two streaks in 
the cell and one in submediaii fold white. Hind wing white 
faintly tinged Avith ochieous. 

Sierra Leone [Clements), 1 J type. Exp. 24 mm. 



128 SIR GEORGE HAMPSON ON THE 

(7) fEMMALOCERA DEPRESSELLA. 

Melissohlajites de2)ressella Swinli. P. Z. S. 1885, p. 876, pi. 57. 
f. 5 ; Hmpsn. Moths lud. iv. p. 63 ; id. Rom. Mem. viii. p. 324, 
pi. 36. f. 12. 

Poli/ocha sacchareUa Dudgeon, J. Bomb. Nat. Hist. Soc. xvi. 
p. 405 (1905). 

Aden ; Puxjab ; Bengal ; Bombay. The lavva feed on the 
roots of sugar-cane. 

(8) tE.MMALOCERA STRIGICOSTELLA. 

Polyocha sirigicosteUa Hmpsn. P. Z. S. 1896, p. 270 ; id. Rom. 
Mem. viii. p. 315, pi. 31. f. 18. 

Aden. 

B. (Emmalocera). Anteuuse of male with short iiuiseriate branches. 

(9) fEjIMALOCERA LEUCOCINCXA. 

Cramhiis leitcocinctus Wlk. xxvii. 169 (1863); Hmpsn. Rom. 
Mem. viii. p. 316, pi. 36. f. 9. 

Emraalocera a^enateUa Rag. Nouv. Gen. p. 38 (1888). 
Singapore ; Borneo ; Philippines. 

C. (Papua). Anteiinx of male laminate, serrate towards base. 

(10) tEiMMALOCEIlA SANGUIFUSALIS. 

Polyocha saiiguifusalis Hmpsn. P. Z. S. 1910, p. 493, pi. xl. 
f. 9. 

X. Rhodesia. 

vll) t-EMMALOCERA AURIFUSELLA. 

Cramhus aurifuseUus Wlk. xxxv. 1756 (1866); Hmpsn. 
Moths Ind. iv. p." 62 ; id. Rom. Mem. viii. p. 317, pi. 36. f. 18. 
Kashmir ; Punjab ; Bombay ; Madras. 

(12) Emmalocera bifidella. 

Polyocha bifidella Wileman, Trans. Ent. Soc. 1911, p. 357, 
pi. 31. f. 22. 
Japan. 

(13) tEMMALOCERA POLYCHROELLA, Sp. n. 

Head and thorax ochreous mixed with fieiy red ; abdomen 
ochreous ; antennas of male with the tuft blackish on inner side ; 
palpi, pectus, legs, and ventral surface of abdomen ochreous 
tinged with red-brown. Fore wing with narrow creamy white 
costal fascia leaving the costal edge red-brown towards base, 
defined below by a fiery rufous streak to beyond middle, the area 
below it red-brown to median nervure and vein 4 ; the inner 



PYRALID.1::. SUBFAMILY HYFSOTKOPIX-E. 129 

half of wing yellow thickly irrorated with fiery i^ed, the terminal 
half of inner margin tinged with brown. 

W. Afuica. (Dudgeon), 1 (5,1 $ ; S. Xigeria, Mama {Dudgeon). 
1 6 type. Ecy)., d ■22, ? 28 mm. 

(14) Emmalocera umbricostella. 

E mmalocera umbricostella Rag. Xou\'. Gen. p. 38 (1888) ; id. 
Rom. Mem. viii. p. 316, pi. 36. f. 10; Hmpsn. Moths Ind. iv. 
p. 62. 

CoREA ; C. Chixa ; SiKHiM ; BoRXEO. Pulo Laut ; Philippixes ; 
Java ; Flores ; Bali. 

(15) Emmalocera lucidicostella. 

Emmalocera lucidicostella Rag. Nouv. Gen. p. 38 (1888); id. 
Rom. Mem. viii. p. 316, pi. 35. f. 20 ; Hmpsn. Moths Ind. iv. p. 62. 
PuxjAB ; Bengal ; Ceylox ; Sumatra. 

(16) Emmalocera anerastica. 

Xepho2iteryx anerastica Snell. V^eth's Midden-Sumatra. Lep. 
p. 81 (1880); Rag. Rom. Me'm. viii. p. 317, pi. 36. f. 8. 

Sierra Leoxe ; Formosa ; Puxjab • oS'icobars ; Johore : 
Selaxgor ; SixGAPORE ; Sumatra ; Borneo, Pulo Laut ; Philip- 
pines ; LouisiADE Is., St. Aignan I. 

(17) Emmalocera latilimbella. 

Papua latilimhella Rag. Bull. Soc. Ent. Fr. 1889, p. ecxx ; id. 
Rom. Mem. viii. p. 313, pi. 36. f. 7. 

Polyocha rhahdota Turner, Pr. R. Soc. Queensl. xviii. p. 122 
(1903). 

Polyocha achrosta Turner. Pr. R. Soc. Queensl. xviii. p. 122 
(1903). 

X. Guinea ; Queensland. 

D. Antenuse of male laminate, not serrate towards base. 

(18) Emmalocera laminella. 

Emmalocera laminella Hmpsn. Rom. Mem. viii. p. 318. pi. 51. 
f. 8(1901). 

Sierra Leone ; Br. E. Africa ; Br. C. Africa. 

Sect. II. Palpi of male dowucurved ; maxillary palpi filiform. 

A. Antenuse of male with sinus and ridare of scales at base of shaft. 
a. Antennae of male with long uuiseriate branches. 

(19) Emmalocera variegatella. 

Polyocha variegatella Rag'. Nouv. Gen. p. 39 (1888); id. Rom. 
Mem. viii. p. 326, pi. 36. f. 16 ; Hmpsn. Moths Ind. iv. p. 63. 

Punjab. 
Proc. Zool. Soc— 1918, No. IX. 9 



130 ■ SIR GEORGE IIAMPSON ON THE 

(20) fEMMALOCERA TRICOLORALIS. 

Polyocha variegatella Hmpsn. J. Bomb. "Nat. Soc. Hist, xii 
p. 320 (nee Rag.). 

Polyocha tricolorcdis Hmpsn. J. Bomb. Nat. Hist. Soc. xv. 
p. 20 (1903). 

SiKKiM ; Philippines. 

(21) Emmalocera diversella. 

Polyocha diversella Hmpsn. J. Bomb. Nat. Hist. iSoc. xii. 
p. 310 (1898) ; Rag. Rom. Mem. viii. p. 324, pi. vi. f. 21. 
Madras, Nilgiris. 

(22) *Emmalocera costella. 

Polyocha costella Rag. Nov. Gen. p. 39 (1888); id. Rom. Mem. 
viii. p. 326, pi. 36. f. 15. 
Gambia. 

(23) Emmalocera cremoricosta. 

Polyocha cremoricosta Rag. Bull. Soc. Ent. Fr. 1895, p. cii ; id. 
Rom. Mem. viii. p. 325, pi. 51. f. 9; Stand. Cat. Lep. pal. ii. 
p. 13. 

Asia Minor ; Syria. 

(24) femmalocera erythrinella. 

Polyocha erythrinella Ra.g. Nouv. Gen. p. 38 (1888); id. Rom. 
Mem. viii. p. 323, pi. 44. f. 14. 

N. Nigeria ; Abyssinia ; Br. E. Africa ; Br. C. Africa. 

(25) *Emmalocera carnatella. 

Polyocha carnatella Rag. Nouv. Gen. p. 39 (1888) ; id. Rom. 
Mem. viii. p. 325, pi. 35. f. 25 ; Hmpsn. Moths- Ind. iv. p. 63, 
Punjab. 

(26) *Emmalocera monochromella. 

Polyocha monochromella Rag. Nouv. Gen. p. 39 (1888); id. 
Rom. Mem. viii. p. 325, pi. 36. f. 13; Stand. Cat. Lep. pal. ii. 
p. 13. 

(27) fEMMALOCERA EREMOCHROA, Sp. n. 

5 . Head and thorax whitish suffused with ochreous brown ; 
abdomen creamy white, dorsally fulvous 3'ellow towards base ; 
})ectus, legs, and ventral surface of abdomen white tinged with 
Isrown. Fore wing pale ochreous, the costal half and terminal 
area suffused with brow n, the cell and veins beyond it with 
some whitish irroration, the interspaces beyond the cell with 



PYRALID^, SUBFAMILY HYPSOTROPINyE. 131 

slight ochreons streaks, the area below the cell with slight red- 
brown irroration. Hind Aving creamy white. 

W. Australia, Sherlock R. [Clements), 1 § type. Exp. 44 mm, 

h. (Lodiana). Antennae of male serrate. 

(28) Emmalocera umbrivittella. 

Lodiana umhrivittella Rag. Nouv. Gen. p. 38 (1888); id. Rom. 
Mem. viii. p. 319, pi. 35. f. 19; Hmpsn. Moths Ind. iv. p. 62. 
Folyocha venosella Wileman, Trans. Ent. Soc. 1911, p. 357. 
Japan, Yezo ; Punjab ; Sikkim ; Assam. 

(29) Emmalocera albicostalis. 

Lodiana albicostalis Hmpsn. Trans. Ent. Soc. 1900, p. 375 ; 
Stand. Cat. Lep. pal. ii. p. 13. 
Palestine; Punjab. 

B. Antennae of male without sinus and ridge of scales. 

(30) Emmalocera subfasciatella. 

Folyocha subfasciatella Rag. Ann. Soc. Ent, Fr. 1887, p. 258 ; 
id. Rom. Mem. viii. p. 328, pi. 36. f. 17 ; Stand. Cat. Lep. pal. ii. 
p. 13. 

Armenia; Persia. 

Auctorum. 

Folyocha rhodemce Strand, Archiv. f. Naturg. 75. 1,3, p. 384 
(1909) N, Rhodesia. 

Genera auctorum. 

Barberia affinitella Dyar, Proc, Ent, Soc, Wash, vii, p. 39 

(1905), nr. Rhinaphe U,S,A,, Texas, 

Cahnia myronella Dyar, J. N. Y. Ent. Soc. xii, p, 108 (1904), 

nr. Saluria U.S.A., Washington, 

Fondotikia translucidella Chretien, Bull. Soc, Ent, Fr, 1911, 

p, 1 1 , nr, Dembea Algeria, 

Schenectadia merilesella Dyar, Pr, U.S. Nat. Mus. xlvii. 

p. 349 (1913), nr. Tinerastia Panama. 

Sabormania pia Strand, Arch. Naturg. 78. A. Hft. 12, p. 80 

1913. '^. = Mo7ioctenoce7'a Spanish Guinea. 



ON VISIBLE AXD INVISIBLE CHARACTERS IN SILKWORMS. 133 

6. First Report on the Inheritance of Visible and Invisible 
Characters in Silkworms. By Miss Maude L. Cleg- 
horn, F.Z.S., F.L.S., F.E.S. ' 

[Received December 3, 1917 : Read March 9, 1918. J* 

The mulberry silkworm races of Bengal are all with the excep- 
tion of one {Bomhyx textor) multivoltine, but their cocoons are 
not so good as those of the European or Japanese races, and 
therefore when the question of reviving the Indian silk industry 
arose, the following suggestions were put forward, viz.: that 
endeavours should be made to obtain an improved multivoltine 
race by crossing an indigenous variety with a European one, and 
that our knowledge of hybridization according to Mendel's law 
should be utilised to help in the process. 

With these suggestions in my mind I obtained from Italy, in 
December 1910, some European silkworm seed (eggs of the 
^ombyx mori) and some Nistri {Bomhyx ct'oesi) seed cocoons from 
the Berhampore Government Nursery. 

The European seed was of the Italian -Japanese Hybrid, and 
was the first generation of a cross between the univoltine (pro- 
ducing one brood a year) yellow Italian (Indigene giallo) male, 
and the univoltine white Japanese female — pure yellow Italian 
seed not having been available at the time. This Italian-Japanese 
Hybrid is very hardy, being a first cross, and so I started my 
cross-breeding experiments with it. 

The cocoons of the multivoltine Nistri silkworm weighed (with 
the chrysalis removed) from 1 grain to r6 grains, whilst the 
cocoons of the Italian- Japanese Hybrid (being of a univoltine race 
and therefore bigger) weighed from about 2*5 grains to 4*6 grains. 
My aim in my experiments with these two varieties of silkworms 
has been (1) to make a multivoltine race (because though its 
cocoons are smaller than those of the univoltine, yet they are 
compensated for by the numerous broods produced during the 
year), producing cocoons of about 4 grains in weight, and (2) to 
see how far the good qualities of the univoltine vai'ieties can be 
combined with the multivoltine character. 

I made two series of experiments : — 

A. A cross between the multivoltine Nistri $ and the univoltine 

Italian- Japanese Hybrid S • 

B. A cross between the univoltine Italian-Japanese Hybrid $ 

and multivoltine Nistri c? . 

Experiment A was made with individuals selected usually from 
three or four families in each generation, except in one of the 
experiments in Fg, which was made with a whole family. 

Experiment B was made with the complete family in each 
generation, with the exception of F,, when only a few worms 
were reared out of four layings. 

The layings produced in Fj by Experiment A and those 



134 MISS MAUDE L. CLEGHORN OX THE INHERITANCE OF 

produced by Experiment B were entirely different from each 
other, for they resembled the maternal parent in each 
case, all the layings of the Nistri g and Ital.-Jap. c5' being 
multivoltine, and all those of the Ital.-Jap. § and Nistri 
c? proving univoltine. 

Result of the Experiment as regards the Size of the Cocoon. 

Experiment A. — The parent cocoons of the Ital.-Jap. S 
and Nistri 5 cross weighed 2*9 and 1-5 grains respectively. The 
cocoons of the first generation of this cross resembled those of the 
Nistri {i. e. the maternal parent) more than those of the Ital.- 
Jap. parent, being rounded at both ends and very thick but 
of the loose texture of the Mstri, the firm texture of the Ital.- 
Jap. being entirely unrepresented. They were uniform in 
size and shape, but varied in weight from 2*5 to 4'1 grains. 

In the second and the immediately succeeding generations, the 
cocoons were not so uniform in size and shape. Many were large 
and rather pointed at the ends, whilst they were all thinner and 
firmer than those of F^, and I found that in the earlier generations 
moths which gave complete multivoltine layings had nearly 
always cut out from cocoons which were about 3 grains or less in 
weight. I did not make use of these 3-grain cocoons for rearing 
purposes, even though they were far better than the original 
multivoltine Nistri cocoon, but I carefully selected the best out 
of the most multivoltine layings of cocoons weighing from about 
3'5 grains to 5 grains in weight. The cocoons in all the generations 
of the cross were far superior to the original Nistri cocoon, and in 
many of the generations they are also sujperior to that of the 
Ital.-Jap. Hybrid. 

Up to Fg many of the cocoons were 4 grains in weight, in Fg, 
F^, Fj, many were over 4*5 grains, while some weighed 5 grains 
and over. In F^ and Fg there were no 4-grain cocoons, and on 
the whole the cocoons of these two generations were very poor 
compared with those of the preceding generations, but they were 
nevertheless still superior to the original Nistri cocoon. As the 
layings from which these cocoons were produced were nearly 
entirely multivoltine, the cocoons appeared to be also becoming 
more multivoltine in character. The rearings in F^ from which 
Fg hatched were all entirely multivoltine, while the Fg silkworm 
were better again and seriposited cocoons, nearly all of which 
were over 3*5 grains, many over 4 gi-ains, and some nearly 5 grains 
in weight (with chrysalis and outer fluff removed). The layings 
of the moths from these cocoons were also entirely multivoltine, 
which showed that it is possible to obtain a 4- or 5-grain cocoon 
from entirely multivoltine layings in Fg after a cross without any 
recrossing. Care was, however, always taken to have the parent 
moths as distantly related as possible. (Table 1.) 

In this expei"iment I found that, after the direct influence of 
the fresh cross seemed to have disappeared, every third generation 
produced the best cocoons, for the cocoons of Fg, F^, F^.^, F^^ 



visible and invisible characters in silkworms. 135 

Table 1, showing the descent of cocoons up to the ninth 
generation, with dates of elevages. 

Ital.-Jap. Hyb.(?yNistri? Jan. 31, 1911 P, 

No. 1, 2-9 grs. ^No. Af^ 1-5 grs. 



No.22? No.l5(? No.26?yNo.l3<? March 28 „ Fi 

3-7 grs. ^3-8 grs. 3-3 grs. ^2-8 grs. 



No.26(? No.33? April 28 „ J'.-, 

4-2 grs. ^ 3-7 grs. 



No.43? No. 21 (? No. 20? June 6 „ F^ 

4"1 grs. 5'1 grs. 5 grs. 



No.59<?' No.l5$ No.5?^No.4(? July 8 „ F^ 

3-8 grs. ^ 4-3 grs. 4*8 grs. ^ 5 grs. 



No.52<? No.19 No. 25 $ No. 18 c? No. 15 ? ^ No. 50 (? No.52J... Aug. 
4 grs. ■^3'8 grs. 4-4 grs. •^3-4gr3. 3'6 grs. 3'2 grs. 4 grs. 



30 „ F. 



No.l0(? No.5? No.l<? No.l7? ^No.l,? Oct. 10 „ F^ 

2-2 grs. ^ 2-8 grs. 3-6 grs. 3-6 grs. ^ 3-6 grs. 



No.41(? No.55? No.61? No.27<? No.41(? Nov. 30 „ F^ 

2-5 grs. ^ 2-4 grs. I'Q grs. ^ 2-9 grs. 2-5 grs. 



No.40?yNo.37<? No.23<?yNo.20? Feb. 1, 1912 i^'g 

3 grs. ^2-8 grs. 2-2 grs. '^2*4 grs. 



No. 115 No. 113 No. 119 No. 221 No. 216 ivlar. 22 „ Fg 

3"7 grs. 4"8 grs. 4"1 grs. 4 grs. 3"3 grs. 



were much superior to those of the intervening generations. 
Even those of F^., seriposited during the rains in July were 
much better than those of F^^ (April) and F^^ seriposited in 
October. As the raiyats in India grow about three or four crops 
of cocoons a year (only rearing a few to keep the breed alive 
when they are short of leaf) it would be to their advantage to 
arrange that the crops reared are of every third generation. 

The cocoons obtained from this multivoltine race are much 
superior to the "Nistri ; those of Fg were valued at about 9J francs 
per kilo in the Milan market, which is close up to the price of 
good Italian cocoons, and the cocoons of F^g, which were not 
near as good as those of F^, were valued at about 7 francs per 
kilo. The correct rendement could not be obtained, as the 175 
cocoons sent to be tested were too few, 1 lb. weight of cocoons 
being necessary. 



136 MISS MAUDE L. CLEGHORK ON THE INHERITANCE OF 

Experiment B. — Cross between univoltine Ital.-Jap. Hybrid 
$ and mnltivoltine Nistri J . 

This was carried out for four generations only and then 
discontinued, as the cocoons were not as good as those of the 
reciprocal cross, rarely averaging a weight of 3'5 grains. 

Residts of Experiment as regards the MtdtivoUine and Univoltine 
character of the Silkivorm Moths. 

Experiment A. 

Table 2, illustrating the descent of the univoltine character in 

THE CROSS BETWEEN ItAL.- JaP. S AND ISTlSTRI $ . 
Multivoltine Nisti-i ? X Univoltine Ital.-Jap. <J ... „ Pi 

Laj'ings all mnltivoltine F\ 

Many more ? s than $ s. 



22 layings univoltine. 30 layings almost univoltine. 6 half multivoltine ^2 

More (? s than ? s. 



6 layings univoltine. 16 about half univoltine. 2 multivoltine JFs 

More ? s than ^ s. 



A few layings univoltine. One 75% multivoltine. Many partly multivoltine ... Fa 
More $s than $s. 



One 75 % multivoltine. 2 half multivoltine. 15 partly multivoltine F^ 

(?s and $ s about equal. 

All about half multivoltine : Jg 



Nearly all about half multivoltine. A few about 75% multivoltine F-; 

Nearly all completely multivoltine F^ 

All multivoltine Fg 

All multivoltine ...........'......;.;...... .Fin 



VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 137 

From the results obtained it will be seen that all the eggs in 
the layings of the multivoltine Nistri $ when crossed with the 
univoltine Ital.-Jap. J hatched, i.e., the layings were all 
multivoltine, whilst the imivoltine paternal parent had no visible 
effect on the character of the layings. This showed that the 
multivoltine character was dominant in the $ , but recessive in 
the (S . Toyama, the Japanese authority on silkworms, also 
found that in crossing pure breeds the first cross always resembled 
the maternal parent. 

I usually found that the completely multivoltine layings did 
not give the most multivoltine results as might be expected, but 
the half multivoltine laying produced the least number of 
univoltine layings in the long run. 

In the Nistri $ and Ital.-Jap. (S cross I discarded- the 
multivoltine layings which appeared in the earlier generations, 
and mostly selected from the half, or partly multivoltine layings, 
till in Fg when most of the layings were almost entirely multi- 
voltine. Some of the families have been entirely multivoltine 
since F^, others have had a few partly univoltine layings, but in 
none of the generations since F^ have any of the layings been 
entirely univoltine. 

Toyama does not give any detailed account of the results he 
obtained in th6 brood characters. It would have been interesting 
and useful, as the inheritance of these invisible characters appears 
to be complicated. He selected from multivoltine forms, but 
does not say if they were entirely or partly multivoltine. He 
states as follows : — " Those forms raised from the first cross do 
not remain true to-the parents in subsequent generations. Even 
when we selected multivoltine forms for five generations we failed 
to get any constant multivoltine breed." 

In his interesting pamphlet "Sulla riproduzione degli Incroci" 
Dott. Quajat, when referring to his experiments with bivoltine 
and univoltine races, states that " nelle successive riproduzioni, 
bivoltinismo tende a diminuire, ed alcune volte anzi a scomparire 
completamente. Sara oria interessante constatare se le ovature 
ottenute univoltine, abbiano alio stato lateute il bivoltinismo, e 
se questo si potra manifestare in seguito a nuovi incroci o 
spontaneamente . ' ' 

I found that from F^, the layings began to show a great 
tendency to become completely multivoltine, and in those of the 
F^j moths, from which F^g worms hatched, only one laying out of 
the whole generation was half univoltine, all the others were 
completely multivoltine. 

To account for the 22 layings in F, appearing univoltine, the 
univoltine character must have dominated in the maternal parents 
of these layings. "When a female F^ was crossed with a " pure " 
multivoltine Nistri c? only 3 eggs out of 250 hatched, and from 
the results obtained, in the reciprocal cross, it was found that if 
the maternal parent was a pure univoltine none of the eggs 
hatched for about a year, and if she was a pure multivoltine all 



138 MISS MAUDE L. OLEGilORN ON THE IXHERITANCIE Of 

the eggs were multivoltine. As this female of ¥^ had hatched 
from a completely multivoltine laying, it might naturally be 
expected that, when the paternal parent was pure multivoltine all 
the eggs laid would lie multivoltine, but as only a few eggs 
hatched it showed that the maternal parent \\as dominant in the 
univoltine charactei-, and that the dominance of the univoUine 
character was inherited hy the F^ $ from the paternal parent in 
which it tvas not a dominant character. 

It is clear that the female influences the leappeai'ance of the 
character in a dominant form, and shows that the descent is of a 
sex-limited inheritance. 

In my experiments I found that the univoltine or multivoltine 
character of the maternal parent showed itself in the layings and 
not that of the paternal, for the character of the paternal parent 
always appeared to be masked or suppressed. 

Besides the difficulties a sex-limited inheritance presents the 
univoltine and multivoltine characters are not visible in the moths 
but only in their layings, for moths which outwardly resemble the 
univoltine parent may have a multivoltine laying and vice versa. 

When all the moths of a generation are bred inter se the 
character of only the maternal parents can be determined by the 
eggs laid ; but to prove that the males and females of each 
generation are either homozygous or heterozygous, dominant or 
recessive, they have each to be bred with pure univoltines and 
multivoltines. So to find out the exact composition of all the 
moths in each generation would requii^e a multitude of experiments, 
and I could not spai-e many moths from F^, for I knew that in 
F., only a very small percentage of eggs would hatch. 

The layings in F3 which were laid by the F,, moths give a clue 
to the chai'acter of some of the parent moths in F... For the 
maternal parents of the six univoltine layings (Table 2) must 
have been dominant in the univoltine character, and those of the 
two multivoltine layings, in the multivoltine character, but the 
sixteen 50 per cent, univoltine layings point to the maternal 
parents being heterozygous. 

Experiment B. 

From the results obtained in the Ital.-Jap, $ and Nistri S 
cross (Table 3) it will be seen that although all the layings in F 
were univoltine, yet there were some multivoltine, and partly 
multivoltine layings in F^, F,, and F^ ; and, again, though the 
moths of F,, were reaied from one of the multivoltine layings 
of F.,, yet 37 layings of the F., moths were univoltine. 

Foi-ty-seven moths from an F,^ family of the Ital.-Jap. $ and 
Nistri S cross were tested with pure multivoltine JSTistri moths 
and the results obtained were as follows : — 

(1) Out of 21 layings of the F,, $ s and pure multivoltine 
Nistri c? s 19 layings were entirely univoltine and 2 entirely 
multivoltine. 



VISIBLE AND TNVISIBLK CHARACTERS IN^ SILKWORMS. 139 

(2) Out of 26 la.yings of pure mnltivoltine Nistri 2 s and F., c? s 
all the eggs liatclierl, thus all being multivoltine. 

These results give a clue to the gametic composition of the F., 
moths, for they show : — 

(1) that 19 1\ $s were dominant in the univoltine character 

and 2 in the multivoltine character. 

(2) that all the E., c? s which may have been univoltine were 

recessive in the univoltine character. 

Table 3, illustrating descent of the univoltine and multivoltine 

CHARACTER IN THE CROSS BETWEEN THE UnIVOLTINE ItAL.-Jap. $ 

AND Multivoltine Nistri cJ • 



Univoltine Ital.-Jap. $ X Multivoltine Nistri (? . 



Layings aJl univoltine Fi 

11 ? motlis 21 (? moths. 



7 layings univoltine. 2 multivoltine. 1 half multivoltine F^ 

62?s.34<?s. 



37 layings univoltine. 2 almost univoltine. 2 multivoltine. 2 almost multivoltine. F3 

34 ? s. 47 (? s. 



14 layings univoltine. 7 almost univoltine. 4 almost multivoltine F4 

(Discontinued.) 

Unequal Sex-ratios. 

I had noticed that in the eai'lier generations of the Nistri $ 
and Ital.-Jap. S cross (which were reared a year previous to that 
of the Ital.-Jap. 5 and Nistri J cross), the number of males 
and females seemed very unequal, and that in one generation 
males predominated, and in another, females. However, these 
remarkable fluctuations graduall}^ decreased in the latter 
generations. 

In my second series of experiments, which were with the Ital.- 
Jap, 5 ^nd Nistri c? ; I noted the exact number of males and 
females in each generation, and found that in this reciprocal cross 
the sex-ratios were also very unequal, but just the reverse in the 
character of the predominating sex, for in F^ of the Nistri 5 
cross there were many more females than males, while in F, of 
the Ital.-Jap. $ cross there were more males than females. 

The unequal sex-ratios could be accounted for by supposing 



140 MISS MAUDE L. CLEGHOR>f ON THE IXHERITANCE OP 

that in some of the generations the univoltine females do not 
hatch, and in others the univoltine, males, but in F^ when all the 
eggs hatched the sex-ratios were also unequal. 

Method of Rearing of Worms. 

To ensure choosing distantly related moths for rearing from, 
I reared the silkworms of each laying seiDarately, and the cocoons 
were also kept separate by keeping the silkworms after their fourth 
moult in rounded trays with a double inner circle of plaited strips of 
bamboo, about an inch and a half from the outer edge, forming 
a space into which the silkworms readily crawl to sereposit their 
cocoons. When the cocoons are removed from the trays, they 
are placed in rows on large sheets of white paper on which is 
noted the number of the laying from which they were reared. 

Each cocoon is covered over with a small earthen cup, the- 
common Indian chilum. A day or two before the moths come out, 
the cocoon is cut open to remove chrysalis and weighed. 

The weight is noted and the chrysalis put back. When the 
moths emerge, the pierced cocoons are removed from under the 
earthen covers and placed within the opening at the top of the 
covers for reference, and this also to prevent the moths from 
getting out. By referring to the number of the layings and to the 
numbers of the two parent cocoons (which is also noted on 
the paper), the choosing of the distantly related moths for 
rearing is simplified. 

If I had been rearing on a large scale I should have kept four 
or six families separate for seven or eight generations, then bred 
them together, and from these again four or six families would 
have been separated out. 

In these experiments the silkworms have always been fed most 
during the night, as I have noticed that, in the wild state, the 
larvae of moths mostly feed up to 9 or 10 a.m., and then restart 
feeding at about 5 or 6 p.m. Even young silkworms, which 
I kept on a small potted mulberry plant under observation, hardly 
ate at all during the day, though surrounded by fresh leaf on all 
sides. I find that silkworms always eat most voraciously between 
10 p.m. and 4 a.m., so it is unnatural for them to be forced to 
feed all day, which is usually the case by the native rearers, and 
may well be one of the causes of their degeneracy. 

They should have a rest of at least seven or eight hours during 
the day, and be fed every three or four hours during the night, 
stai^ting late in the afternoon at about 6 p.m. If there is an 
abundant supply of leaf, they might with advantage be fed every 
two hours. Newly hatched worms should always be fed every 
two hours between 5 p.m. and 10 a.m. The worms if fed well in 
the last stage after the fourth moult will give good cocoons, even 
if not very well fed in the earlier stages, but it is most essential 
that they be well fed throughout the larval stage for them to be 
vigorous and healthy. 



VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 141 

One of the advantages gained by the nocturnal feeding of 
worms is that the leaf can be gathei-ed at sundown, and so keeps 
fresher, and does not fade as readily as w-hen plucked in the 
morning. There is also very little chance of attacks from the 
dreaded silkworm fly, which is always on the alert during the 
brightest and hottest hours of the day. 

I have found that the leaf from male trees of Morus Indica 
gives better results in feeding silkworms than that of the female 
trees. The dioecious character of the midberry is not, as far as I 
know, known to sericulturists, for in most handbooks on the 
subject they advise rearers to select trees which bear little or no 
fruit. The inconspicuousness of the catkins and pseudo-spikes of 
the mulberry has evidently led to the difference between the 
trees being overlooked by all but botanists. As the flowers are 
inconspicuovTS they are anemophilous, and the trees, when in 
flower in February, may be seen giving off pollen in little puffs 
like smoke. This is best observed when there is no breeze. 

Additional Observations made during Expeonments. 

The inheritance of the visible colour character of the cocoons is 
clearly Mendelian. 

The parent cocoons of the Ital.- Jap. Hyb. were of the yellow 
Italian cJ and white Japanese 5 . 

The colour of the yellow Italian cocoon is of a deep pinkish 
yellow {carneo-giallo or flesh colour), which sometimes varies from 
a deep orange to almost white, but never of the vivid yellow of 
the Indian cocoon. All the cocoons of F^ of this hybrid were of 
a pale flesh colour, none were white like that of the Japanese 
parent. A character which dominates after a cross is made was 
described by Mendel as dominant, and the character which 
seemed to have disappeared he called recessive. So the flesh 
colour of a cocoon is a dominant character, and the white 
a recessive. All observed results in the study of heredity point 
to the dominance of a character being due to the presence of that 
character, and the recessive to the absence of the dominant 
character. 

The flesh colour of the Italian is due to the presence of the 
flesh colour and to the absence of the vivid yellow colour of the 
Nistri. While the white colour of the Japanese cocoon is due to 
the absence of both the pinkish yellow and bright yellow colours 
— the example of the inheritance of eye-colour in man might 
make this clearer. Here the dominance of brotvn eyes can be 
traced to the presence of a brown pigment ; and the recesslveness of 
blue eyes to the absence of the broivn pigment. All human eyes 
(except those of Albinos) have a layer of deep purple pigment on 
the inner surface of the iris, but in brown, hazel, green, and giey 
eyes there is also a layer of brown on the outer surface of the iris, 
and it is this brown layer which entirely (if abundant) or partly 
conceals the purple layer. In clear grey and clear deep and pale 



142 MISS MAUDE L, CLEQHORN ON THE INHERITANCE OF 

blue eyes the brown pigment is absent and the purple pigment 
shows through the tissue forming the iris, and makes it appear 
of a clear deep blue, when the tissue through which it is seen is 
very delicate — and of a clear grey or pale blue when the tissvies 
are more or less coarse. So, to trace the dominance of brown 
eyes the real nature of the variovis kiuds of light eyes must be 
carefully made out; it then becomes clear that the presence of the 
hrown pigment in brown eyes makes broivn eyes dominant to blue 
eyes in ivhich it is absent. If one parent has' very dark brown 
eyes and the other clear blue eyes, all the children willhave dark 
eyes, which sometimes include hazel, green or grey, but none will 
have clear grey or blue eyes. The reason for there being no 
blue-eyed children is apparent, for all the children inherit a 
factor, or unit character for pigment from the dark-eyed parent, 
but a factor lacking in j^igment from the blue-ej^ed parent — and 
the presence of this pigment in all the children makes brown eyes 
dominant to blue. 

To return to the colour of the cocoons the simplest explanation 
will be found in the interaction of two simple Mendelian 
characters. These two characters are flesh colour F, and yellow 
colour Y, and the two pairs of unit characters involved are — 

1. Flesh colour F. Absence of flesh colour f. 

2. Yellow colour Y. Absence of yellow colour y. 

The parent moths, then, have the following constitution : — 

(1), Yellow Italian possessing the flesh colour and lacking the 
yellow colour of the Nistri ... ... ... FFyy 

(2) Japanese white lacking both characters ... . . ffyy 

(3) Nistri lacking the flesh colour and possessing the yellow 

colour YYff 

The actual parents in the experiment were : — 

P^ the Ital.-Jap. Hyb. an F^ of a cross between the Yellow 
Italian and White Japanese. 

P, the ISTistri. 

The Yellow Italian has a deep pinkish yellow (flesh coloured) 
cocoon. It inherits two factors, one for the flesh colour F, anel 
oiie for the absence of the yellow colour y, from each parent, and 
so consists of the union of two similar pairs of factors Fy and Fy. 
The Yellow Italian is therefore pure (homozygous) as regards the 
colour, for the germ-cells (gametes) by the union of which it was 
formed, each carried the same kinds of factors Fy and Fy. So 
the gametic composition of the Yellow Italian for the colour of its 
cocoon is represented as FFyy in Table 4, Diagrams (1) and (2). 

As the whiteness of the Japanese cocoon is due to the absence 
of both the flesh colour of the Italian, and the yellow of the 



VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 143 

Nistri, the gametic composition is represented in small letters as 
ffyy (Table 4). There was no colour in either of its parents for 
it to inherit, for the gametes, from the union of which it was 
formed, bore the factors fy and fy. Table 4, Diagram (1). 

The cocoons of the Ital.-Jap. Hyb. are a pale flesh colour as 
they inherited the flesh colour from the Yellow Italian parent 
only, for the gamete from the White Japanese parent bore colour 
factors lacking in both the flesh and yellow colours. The colour 
factors inherited from the Yellow Italian parent were ^y 
and from the white Japanese parent fy. The Ital.-Jap. Hyb. is 
therefore not pure as regards the colour of its cocoon, for the two 
gametes, from the union of which it was formed, were unlike and 
did not carry pairs of similar factors. It is a hybrid (heterozygote), 
the gametic composition being Ffyy, and it will give gametes Fy 
and fy. When the Nistri (YVff), Avhich has a deep yellow 
coloured cocoon, was crossed with the Ital.-Jap. Hyb. all the 
cocoons of Fj of this cross were of a bright yellow colour, but not 
quite the deep yellow of the Nistri. So the bright yellow of the 
Nistri was dominant to the pale flesh colour of the Ital.-Jap. Hyb., 
for the deeper yellow entirely masked the pale pinkish yellow 
, of the flesh colour even when it was present (Table 4). The 
gametic composition of F^ of the Nistri $ and Ital.-Jap. (S cross 
was YyFf and Yyff, and it therefore contained two classes, 
both numei"ically equally represented. Table 4, Diagram (1).' 
All the cocoons were yellow, but not quite uniform in tin. 

The gametes given ofi' by these two classes were : — 

YyFf giving YF, Yf , Fy, yf. 
Yyff „ -Tf, yf. 

In F., there were a few deep yellow cocoons like those of the 
Nistri, a good many bright yellow like those of F^ a few pinkish 
yellow and a very few white out of 52 cocoons. The exact 
numbers were — 13 deep yellow, 29 yellow, 6 flesh coloured, and 4 
white out of 52 cocoons. 

The gametes for the colour character in F^ were YF, Yf, Fv, 
and yf, and their union at random would give, in F.,, cocoons of 
the following gametic compositions — YYFF, YYFf, YYff, YybF, 
YyFf, Yyff, yyFF, yyFf, and yyff, which i^esulted in cocoons of 
various shades with just a few white ones. 

This is just what occurred, and it will be seen that the 
proportions of the various colours obtained out of 52 cocoons 
run the proportion calculated very close, for according to the 
analysis in Table 4 there should be 12 deep yellow, 24 yellow, 
5 flesh coloured, and 7 white out of every 48 cocoons. Table 4, 
Diagram (2). There was a difliculty, however, in distinguishing 
between the deep yellow and the ' blight yellow from the 
intermediate forms which occurred. 

In F3 cocoons I found that in some cases when both the jjai-ent 
cocoons were deep yellow, all the cocoons produced by the ofispi'ing 



1 44 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OP 

were deep yellow too ; in other cases, two deepish yellow F^ 
cocoons produced some flesh-coloured and even white ones. 

I also found that flesh-coloured parent cocoons would, in some 
cases, give only flesh-coloured ones, and in others flesh-coloured 
and white. 

Table 2 shows that among all the yellows in ~F^ there are two 
kinds — the pure (homozygous) dominants YYff and YYFF and 
the hybrid (heterozygous) dominants Yyff , YyFF, and YyFf. So 
when YYff and YYff are chosen as the parents, all the offspring 
will produce deep yellow cocoons, but if YyFF and YyFf are 
chosen, about a quarter of the cocoons will be flesh-coloured, and 
if the light yellow Yyff and Yyff are chosen as the parents, about 
a quarter of the cocoons will be white. 

In his experiments with the white Japanese and yellow 
Siamese, Toyama obtained from the F^ yellows some which 
produce only yellows, and 2 yellow cocoons which gave 221 yellow 
and 77 pinkish yellow, and 2 yellows which produce 254 yellow 
and 77 white cocoons. These results obtained by him give almost 
the exact proportions (75 per cent, and 25 per cent.) I have 
obtained in my analysis of the colour factoi-s, Diagiams (2) 
and (3) Table 4. 

Among the flesh-coloured cocoons they are two kinds — the 
homozygous FFyy and the heterozygous Ffyy. Parent cocoons. 
' which are both FFyy will produce all flesh-coloured ones, but 
two Ffyy parent cocoons will give a few white ones. Table 4, 
Diagram (3). Toyama appears to have obtained both the pure 
and impure flesh-coloured cocoons in his experiments. For, 
referring to them, he states " the pale-pinkish-yellow form pro- 
duces some uniform (producing only pale-pinkish-yellow) and 
some mixed (the white 25 per cent, and the pale-pinkish-yellow 
75 per cent.) offspring in each succeeding generation." 

The pure white cocoon bred inter se always gave white cocoons. 
The yellowish white cocoons. The yellowish-white ones, how- 
ever, mostly gave a small percentage of yellow cocoons. 

In the gametes of the pure white cocoons there is no colour 
factor, so when the parent cocoons are both pure white the 
offspring inherit only colourless factors and will all be white. 
Table 4, Diagram (3). 

The white recessives are also easily distinguished from the 
dominant yellows, and as they are pure (homozygous) in the 
colour character, a race with white cocoons can be easily 
made from white cocoons which appear in any of the generations. 
Toyama also found that " every white form from its first pro- 
duction remains true to itself." 

The recessive character always breeds true in whatever genera- 
tion it occurs, but as it never is present in the F^, after a cross, 
the F individuals must always be reared, as the recessives make 
their first appearance in F^. 

On the whole the results of these experiments, excepting that 



VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 



145 








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Proc. Zool. Soc— 1918, No. X. 















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146 ox VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 

of the visible colour character, show great complications. The 
univoltine layings which appeared in tlie generations after F^ in 
the multivoltine Xistri 2 '-^^^*^ univoltine Ital.-Jap. S cross, and 
the multivoltine layings which appeai'ed in the recipiocal cross, 
show that the maternal parents are dominant in the univoltine 
and multivoltine character respectively, and that these characters 
ivere inherited from the paternal grand-parents in which they luere 
dominant characters. So these recessive characteis in males 
appear to become dominant when inherited by the females. 
Either the female sex is in some way closely associated with the 
dominance of the multivoltine and univoltine characters, or there 
is some factor present in the male which makes the univoltine and 
multivoltine characters lie latent, but does not hinder them from 
being handed down to the offspring, the females of which may show, 
in a dominant form, the latent cbai'acter of their paternal parent. 
The inheritance of the invisible vmivoltine and multivoltine 
character does not appear to be quite Mendelian ; however, it 
may he that the sex-limited descent afiects the inheritance, and 
there is really no failure in the segregation of the unit characters 

Bibliography. 

Bulletin of the College of Agriculture. — Tokyo Imperial Univer- 
sity, Japan, 1906, Vol. VII. "Studies on the Hybridolog}' of 
Insects '' : K. Toyama. 

" Sulla Riproduzione degli Incroci e su aleuni cai-atteri ei-editai-i 
che presenta la, Sericaiia Mori in relazione alle leggi di Mendel '' : 
E. Quajat. 

"Mendel's Principles of Heredity": W. Bateson. 

" Breeding and the Mendelian Discoveiy " : A. D. Darbishire. 



ON CETACEA STRANDED ON THE BRITISH COASTS. 147 

7. Notes on Cetacea stranded on the British Coasts during 
1913-1917. By Sidney F. Harmer, Sc.D., F.R.S., 
F.Z.S., Keeper of Zoology in the British Museum 
(Natural History) *. 

[Received March 19, 1918 : Read March 19, 1918.] 

The stranding of specimens of Cetacea on vai-ious parts of the 
British Coasts is no new thing, but in the majority of cases the 
evidence derivable from these occurrences has not been sufficiently 
used. About six years ago the Trustees of the British .Museum 
decided to examine this evidence more systematically ; and an 
arrangement was accordingly made with the Board of Trade 
and with the Admiralty, by which instructions were issued to 
Receivers of Wreck and members of H.M. Coastguard that the 
stranding of specimens of Whales should be reported by telegram 
t© the Museum. These orders were given during 1912, and the 
system had become fully operative by the beginning of 1913. 
The notes here recorded .give a summary of the results thus 
obtained. 

The telegrams received are based on leaflets which were 
distributed on behalf of the Museum, calling attention to some 
of the more obvious characters by which the species of Cetacea 
can be distinguished. They have generally been sufficient to 
give some preliminary idea of what tlie stranded animal was 
likely to be ; and further evidence has been obtained, wherever 
possible, by means of correspondence and by securing the 
specimen in question, or some part of it, when this could be done. 
A written Keport, on a form requesting answers to certain 
specified questions has generally proved very instructive, 
particularly in cases where the answers have been supplemented 
by sketches or photographs. 

The species of Whalebone Whales can usually be distinguished 
by the characters of their whalebone or baleen ; and a special 
effort has accordingly been made, in each such case, to obtain a 
blade of baleen. In the case of the Toothed Whales, the receipt 
of a lower jaw is often sufficient for specific determination ; but 
in a number of instances the entire head and flippers have been 
secured, or even the entii-e animal or its skeleton. The records 
may accordingly claim to have been based on satisfactory 
evidence, in the majority of instances in which a positive result 
h.a,s been recorded. 

During the five years under review, fifteen species out of some 
twenty generallj^ recognized as British have been recorded. The 
.species which have not at present been reported are the Atlantic 
Right Whale or Nordkaper {Bcdcfna glacialis Bonn.), the Blue 
Whale or Sibbald's Rorqual {Balceno'ptera inv.sculus L.), the 

* Published by permission of the Trustees of the British Museum. 

10* 



148 DR. SIDXEY F. HARMER ON CETACEA 

Humpback {Megcqitera nodosa Bonn.), the White Whale 
[Delpkinapterus leucas Pall.), and the Narwhal [Monodon 
Dionoceros L.). The following observations refer in the first 
instance to specimens actually recorded as having been stiunded 
during the live years in question (Harmer, 1914-1918); though 
the statements of previous observers are noticed to some extent. 

It may be remarked that the number of records of the larger 
Cetacea in the neighbourhood of the British coasts has been 
largely increased during recent years by the I'esults of the 
Whaling Companies which have operated at certain Stations 
in Scotland and Ireland. Information on this subject is given, 
for the Scotch Stations, by Thompson (1912, p. 393), and for 
the Irish Stations in a 23aj)er by Lillie (1910), and in the Reports 
to the British Association by Burfield (1913) and Hamilton 
(1915, 1916); as well as in other memoirs cited by those authors. 



, MYSTACOCETI or WHALEBONE WHALES. 

(1) Lesser Rorqual [Bahenoptera acutorostrata Lacej).). 

This has proved to be the species of Whalebone AVhale most 
commonly stranded on our coasts ; and twelve specimens have 
been defi.nitely recorded (including two from 1911 and 1912 
respectively), besides one or two others whicji probably belonged 
to the same species. All have been found duiing the summer or 
early autumn, from June to October; and from two distinct 
regions of the coast : — (a) the Eastern coast of Scotland and 
E.igland, from Caithness to Yorkshire; (6) the South- Western 
district, including the Xorth coast of Cornwall, the Welsh coast 
and the South- West of Ireland. 

The Lessei' Rorqual reaches a length of about 33 feet, and 
it can be recognised by the broad white band which i-uns across 
the outer side of its pectoral limb or flipper and by the colour of 
its baleen, which is entirely white or yellow, with hairy fringes 
of the same colour. The baleen-blades may, however, have a 
rosy tinge at their base, owing to the presence of vascular papillse 
containing blood. 

Although the British Museum possesses but little material 
bearing on this point, I believe that there is a material difference 
in the thickness of the hairs of the baleen-blades between young 
and fully adult specimens. Thus 1 find that in the baleen of a 
Lesser Rorqual, stianded at Perranporth, Cornwall, on June 5, 
1916, the hairs are much finer than in an individual of the same 
species stranded at Ulronie, Torkshii'e, on Oct. 21, 1915. In 
both instances the determination of the species is confirmed by 
other evidence. Tlius in the specimen fi-om Perranporth the 
pectoral fin was described as having a white band on its outer 
surface — a marked specific character of the Lesser Rorqual ; 
and this evidence was completed by a sketch of the pait in que.s- 
tion. In the animal from Ulrome the baleen was desci-ibed as 



STRANDED ON THE BRITISH COASTS. 149 

being all of one colour, white or j^ellow — another distinguishing 
feature of the same speciey. The Perranportli Whale was only 
18 feet long, while that from Ulrome measured 33. feet. The 
much finer character of the baleen-hairs of the former specimen 
was thus probably due to immaturity. 

This species is known to occur off the Norwegian coast at all 
times in the year, and it feeds to a large extent on fish. 

(2) RuDOLPHi's Rorqual or Sbi Whale (Balceiwptera borealis 
Less.). 

In my Annual Reports on Stranded Oetacea (1914-1918) I have 
recorded the following four specimens as belonging to this 
species : — 

1914, l^eb. 28.— Derrynane, Co. Kerry, 60 ft. 

,, Sept. .21. — Crosskirk, Caithness, 43 ft. 

,, Nov. .17. — John o' Groats, Caithness, 47 ft. 
1917, Oct. 13. — Annet, Scilly Islands, 45 ft. 

These determinations wei'e made mainly on the evidence of 
single blades of baleen ; and I have to admit that on a re- 
examination of these specimens, and taking into consideration 
the other evidence available, I have come to the conclusion that 
they were probably incorrect in three of the four cases. The 
mistakes (if they were wrong conclusions) arose principally from 
the comparison of young specimens with old ones. 

There can be little doubt that the specimen from Crosskirk 
was correctly determined. Its baleen-fringes ai'e composed of 
extremely fine, flexible hairs, which have the curly, -wool-like 
texture which has been noticed by other writeis on Rudolphi's 
Rorqual. In this respect they agree precisely \\itli a series of 
baleen-plates, 1912.6.25.1 in the British Museum collection, 
obtained from one of the Shetland Whaling Companies and 
undoubtedly correctly determined. In the other three specimens 
which were referred to B. borealis, the baleen-hairs, although 
relatively fine, particulai'ly in the individuals from John o' Groats 
and the Scilly Islands respectively, were straight and not cuiIy. 
The fact that they are finer than those of the large blades of a 
fully grown Common Roi'qual is probably due to immaturity*, 
and perhaps to the small size of the blades, which may have been 
taken from near one end of the baleen-series. I am accordingly 
led to the conclusion that the three individuals in question m ere 
Common Rorquals; and some further evidence in support of this 
conclusion is forthcoming. 

It was pointed out by G. 0. Sars (1881, pp. 3, 5) that the 
Common Rorqual is characterized by a very remarkable asymmetry 
in the coloration of the head, and that this featvire is invaiiable 
in both sexes and at all ages, at any rate in the Northern 

* See the remarks on this subject under the lareceding species. 



150 DR. SIDNEY F. HARMER OX CKTACEA 

specimens which were examined by this observer. On the left 
side of the head, the upper jaw and the lo\\'er Hp ai'e dark in 
coloui', like, the doi'sal surface generally. On the right side, the 
anterior end of the upper jaw, the greater p&vt of the lower jaw 
and a number of the anterior baleen-blades are white; these 
blades, as Sai's points out, being much like those of the Lesser 
Roi-qua,l. 

In the specimen from John o' Groats the baleen-plates weie 
described as being blue on one side, and white for a distance of 
two and a half feet from the front end of the beak on the other 
side, the remaining plates of the same side being blue. It is 
fortunate that tv\o photographs of this individual were supplied, 
taken from the two sides. In these, the upper part of the head and 
the lower jaw appenr dark on the left side, while no difference 
in the colour of the baleen is apparent anywhere in the series. 
On the right side, the front part of the upper jaw, the lower lip 
and the baleen-plates of the anterior end of the series are white ; 
and these character's make the reference of the specimen to 
B. phi/scdus practically certain. 

In the individual from the Scilly Islands the evidence is less 
complete, though the baleen in the fore part of the mouth was 
described as having been white, " getting gradually darker till it 
becomes black."' In the photographs received of this individual (see 
p. 13 of my Report on the specimens stranded during 1917), the 
left side is not visible. On the right side, however, the lower 
lip at least is white, and apparently part of the upper region of 
the head. The baleen is iK)t visible. The blade received is a 
small one, probably from near one end of the series. It agrees 
so closely with that of the individual from John o' Groats, in the 
fineness and straight character of the hairs, that I feel little 
doubt that it belongs to the same species ; while it may further 
be remarked that its coloration agrees better with that of the 
baleen in the Common Rorqual than with that in Rudolphi's 
Rorqual. This specimen, too, thus seems to have been an 
immature B. pliyscdus. 

The evidence with regard to the specimen from Derrynane was 
very inadequate ; but I am inclined to refer this one also to 
B. jihysalus on the evidence of a single incomplete baleen-blade, 
in which the hairs are not cui'ly, but straight and somewhat 
coarser than in the other two specimens, in coi-relation with its 
greater size. In my original record of this specimen I remarked, 
on the assumption that it was a Rudolphi's Rorqual, that the 
length (60 ft.) recorded was probably too great; but if the 
determination then made was incorrect there is no reason to 
suppose that the size of the animal was overestimated. 

If the foi'egoitig corrections are justifiable, the only record of 
B. horealis, in these observations, is the specimen from Crosskirk, 
Sept. 1914. According to Gollett (1912, p. 597), this species, 
which feeds principally on pelagic Crustacea, only reaches the 
Norwegian coast during the summer. It becomes from 45 to 50 



STRANDED ON THE BRITISH COASTS. 151 

feet long when adult, and its baleen is usually black in colour, 
though spnie of the blades are pai'tly white, with white hairy 
fringes of the fine, silky texture and curly character noted aloove. 
An elaborate account of this species has I'ecently been given by 
Andrews (1916); while the memoir of Allen (1916, p. 234) may 
also be consulted for its coloiation and other charactei's. The 
maximum length of B. borexdis is said to be about 53 ft. 

(3) Common Rorqual {Balcenoptera physcdus L.). 

Including three of the individuals which weie originally 
referred to B. horealis, as above explained, this species is 
represented by eight records : namely, North Devon (Feb., 1913), 
Kerry (Feb., 1914), Northumberland (May, 1915), Caithness 
(July, 1913), Donegal (Aug., 1913), North Kent (Oct., 1914), 
Scilly Islands (Oct., 1917), and Caithness (Nov., 1914). 

The Common Rorqual usually reaches a length of 65 to 70 feet 
in Northern waters. Its baleen-blades are for the most part 
slate-coloured, with darker and lighter longitudinal streaks; but, 
as pointed out in the account of the preceding species, a number 
of them at the anterior end of the right series are white. Their 
hairy fringes are yellowish in colour. The food consists partly of 
fish and partly of pelagic Crustacea, the species showing a , 
preference for the latter form of diet (Collett, 1912, pp. 581, 582). 
Like the Lesser Rorqual, it occurs of!" the Norwegian coast at all 
times in the year. 

B. physcdus is known to be very variable in colour; and to such 
an extent that various forms of the species have been distinguished 
by the Norwegian whalers. For information on this subject see 
Cocks (1884, p. 458 ; 1887, p. 2L5), True (1904, p. 119), and Allen 
(1916, p. 181). The specimen here recorded from the Scilly 
Islands appeared to have a good deal of white on the dorsal 
surface, a type of coloration which is unusual for this species. 
But I have recently been informed by Mr. King, who took the 
photographs published on p. 13 of my Report for 1917, that in 
his opinion much of the white colour observable was due to the 
loss of the skin in patches and the consequent exposure of the 
underlying blubber. 

ODONTOCETI or TOOTHED WHALES. 

Fam. Physeterid^. 

Subfam. PHYSETERINiE. 

(4) Sperm Whale {Physeter catodon L.). 

Three records, from E. Caithness (May, 1917), Gaiway (Sept., 
1916), and Inverness (Dec, 1913). 

The Sperm Whale, which occurs principally in the warmer 
seas, is remarkable for the striking ditfeience in size between the 



152 DR. SIDNEY F. HARMER ON CETACEA 

males and tlie females; the males reaching a, length of at least 
60 feet, and the females probably not much n.ore than half that 
length. Its most obvious character is the possession of a long 
series of exti;emely large teeth, about 20-25 in number, as much 
as 8| in, in length and 3k or even 3| in. in basal diameter, on 
each side of a narrow lowei- jaw ; the upper jaw being edentulous 
or with vestigial teeth of irregular form. The maxillary teeth of 
the Sperm Whale are figured by Owen (1840-1845, pi. 89. figs. 3, 
4), and by Sir William Turner (1912, pi. ix.), according to whom 
they may be as many as 15 in number, and may reach a length of 
80 mm. (p. 74). Owen (p. 354) gave the number as 8 on each 
side. Other striking features of the species are the enormous 
truncated head extending some distance in front of the tip of the 
lower jaw, and the position of the blow-hole on the left side, at 
the anterior end of the head. 

Many records of the occurrence of the Sperm Whale in the 
Biitish seas have pieviously been published, a large proportion of 
them having been on the Scotch coast (cf. Turner, 1871, 1872, 
1904). Part of the skull of a large Sperm Whale, stranded in 
1 582 at Caister, on the Norfolk coast, may be seen in the Ohui-ch 
of St. Nicholas at Great Yarmouth. Mr. A. B. Van Deinse has 
just published a paper (1918) on 37 specimens recorded on the 
■ Dutch coast during the period 1531 to 1788. It is a remarkable 
fact that, with very rai-e exceptions, the Sperm Whales recorded 
in European waters are of large size and are therefore presvimably 
males. It is believed by the Whalers that these are roving 
individuals which have been driven away from the hei'ds by the 
competition of other males (cf. Thompson, 1912, p. 397); the 
species being polygamous. Although two of the records here 
given conform to the general rule, the thiid, from Galway, is of 
special interest ; having been a young individual, with uncut 
teeth, of only 18 feet in length. The condition of the dentition 
shows that this individual was a "sucker"; and that an adult 
female must have been somewhere in the neighbourhood, although 
its presence was not recorded. 

The Sperm Whale feeds largely on Cuttlefish, but partly at any 
rate on fish. 

Subfam. Ziphiin^, 

The Ziphioid Whales are distinguished from the Physeterinse 
by a further reduction of the dentition ; the functional teeth 
being commonly a single pair, or more rarely two pairs, in the 
lower jaw. Other vestigial teeth have, however, been recorded 
in both jaws in all the three genera known in British seas. The 
functional teeth generally remain beneath the gum in young 
individuals and in the females even when adult; but they pierce 
the gum in males sooner or later, sometimes only when fully 
adult. In Hyperoodon and Ziphius there are, moreover, strongly 
marked differences in cranial characters between the adults of the 
two sexes. 



STRANDED OiN TUE BRITISH COASTS. 153 

(5) BoTTLE-2\osED AV^HALE {Hypevoodon rostratus Miill.). 

Fourteen records, mostly from the Nortliern coast of Scotland, 
along the entire Eastern coast of Scotland and England, and in 
the South of England as far west as Somerset. 

The Bottle-nosed Whale has long been hunted in Northern 
seas (cf. Collett, L906), .and its movements are better known 
than those of manj^ other species. According to Hjort (1912, 
p. 649, chart on p. 650) it is present in considerable numbers in 
the Norwegian Sea during April-July; extending as far North as 
Lat. 76° in June. In September, when it is migrating South, the 
Faroe Islanders get their last Bottle-noses. The localities where 
it is principally found are on the Western side of the Gulf 
Stream water, in the transition-belt between the Arctic and 
Atlantic currents. "The individuals of this species are said to 
follow the 800-fathom line dui-ing their migration. 

It has been pointed out by several authorities (cf. Turner, 
1-886, p. 45) that Buttle-nosed Whales are most commonly 
stranded on the British coasts in the autumn, when on their 
Southward migration ; the individuals recorded being either 
young specimens of either sex or adult females, often accompanied 
by a calf. The occurrence of the species, as shown by the British 
Museum records, is in general agreement with this statement, 
except that the sex of the majority of the specimens is not 
definitely known and there are no records of females with calves. 
There is one record for each of the months July (1917) and 
August (1916), both from Scotland ; four for each of the months 
September and October ; and one from the South of England 
(Somerset) in November, 1914. The only others of which 
evidence has been obtained are two in March (Northumberland, 
1914 ; Caithness, 1915), and one near Nairn, in May, 1914. 

A remarkable altei-ation is knoAvn to occur in the male Bottle- 
nosed Whale, with increasing age, both in external form and in 
cranial characters. The changes in the skull ai'e principally due 
to the immense increase in the height and thickness of the 
maxillary crests which are so characteristic a feature of this 
species, the adult males having formerly been described as a 
distinct species, H. latifrons (xray. With this increase in the 
maxillary crests is associated a corresponding change in the profile 
of the head ; the "forehead" becoming more and more prominent 
until in old age its antei'ior outline is quite vertical. The old 
males appear to have a migration-route which is further from the 
land than that of the adult females ; with the result that they are 
hardly ever recorded on the British coasts (Turner, 1886, p. 45). 

According to Southwell (1883, p. 480) the young Bottle-nose is 
black, the colour becoming lighter with age ; old specimens being 
almost yellow, with a greyish white ventral surface, while 
the beak and front of the head are quite white. The oldest 
males may become white all over (Millais, 1906, p. 298). 

This species is provided with a pair, or sometimes two pairs, of 



154 DR. SIDNEY F. HARMER ON CETAOEA 

large teeth, which may reach a length of 40 mm. (Collett, 1906, 
p. 11), at the extreme front end of the lower jaw. According to 
most authorities these teeth remain concealed throughout life in 
females, but they pierce the gum in the oldest males, and are then 
commonly provided with a tuft of the Stalked Barnacle, 
Conchoderma auriium. Vestigial teeth, in addition to these, 
have frequently been described in both jaws (figuied by Turner, 
1912, p. 82) ; and traces of them have been found in one or two 
of the individuals here lecorded. The niale in this species reaches 
a length of at least 30 feet, while the female is said not to groAv 
much beyond 24 or 25 feet (Munsterhjelm, 1915, p. 9). Collett 
(1906) states that the Bottle-nosed Whale is believed to be able to 
remain under water at least two hours, and that, although it feeds 
principally on Cephalopods, it also eats Herring or Cod, or even 
pelagic Crustacea. Newly-born young have been observed in 
June, while other females possess a small fcetus at this season. 



(6) Cuviee's \^ hale (Zip/lilts cavirostris Cuv.) *. 

Before these records were instituted, this species was known 
as British on the evidence of a single skull, obtained by Sir 
William Turner from the Shetland Islands. One of the most 
interesting results of the system inaugurated by the British 
Museum has been the demonstration that Cuvier's Whale is 
not the extreme rarity it had been supposed to be. 'I hree 
individuals have been definitely determined, from Cork (Feb., 
1913), North Cornwall (June, 1916), and Wexford (July, 1915). 
It is by no means unlikely that some of the older records of 
" Bottle-nosed Whales " may have belonged to this species. 
Two of these specimens have already been recorded by me in a 
communication to this Society (Harmer, 1915, p. 559). 

True (1910, p. 54) has given reasons for believing that the 
adults of the two sexes of this species are distinguished by marked 
dijferences in ciunial characters and in the teeth. In adult males, 
according to this authority, the mesirostx'al ossification is enor- 
mously developed and there is a deep " prenarial basin " in the 
skull. The teeth, of which one pair are present at the front end 
of the lower jaw, as in Hyperoodon, are fusiform and leach a 
diameter of 25 to 30 mm. In adult females, the mesirostral 
ossification is only slightly developed and a prenarial basin is not 
formed. The teeth are slender, with a diameter of 10 to 14 mm. 

Although the skulls of the specimens here recorded are not yet 
all available for study, they appear to confirm True's statements. 
The largest individual (Co. Cork) measured 26 feet in length, 

* [In my Annual Report for 1917, specimen No. 7, recorded on June 9 from 
Co. Clare, was described as a Ziphins. The examination of its skeleton, which has 
just been cleaned, proves that it belongs to a species of Mesoplodon, having its two 
teeth at the extreme anterior end of the lower jaw. It is proposed to publish 
a further account of this highly interesting specimen in due course. — S. F. H., 
July 16, 1918.] 



STRANDED ON THE BRITISH COASTS. 155 

and its cranial charactei'S agree with those regarded by True as 
belonging to adult females. The teeth are slender, with a 
diameter of about 13 mm., and were completely concealed beneath 
the gum. Although the sex was not ascertained from observations 
made on the entire vspecimen, there can be little doubt that 
the animal was an adult female. The Cornish specimen was 
incomplete, the part recorded measuring 15 feet in length. Its 
teeth were uncut, and it was presumably a young male or female. 
The other specimen, 18 ft. 2 in. long, was definitely known to 
be of the male sex. Its teeth were large and massive, with a 
diameter of 35-37 mm., and they projected beyond the gum. It 
thus appears probable that the teeth remain uncut in the female 
Ouvier's Whale throughout life, unless they become external in 
old age, while in the male they are probably cut relatively 
early. 

The coloration of this species appears to be variable. Certain 
individuals have been recorded in which the upper part of the 
head was white; and the Wexford specimen possessed this type 
of coloration. In other cases the colour has been described as 
dark above and lijjht beloAv. 



(7) Sowerby's Whale (Mesojdodon Helens Sowb.). 

This rare species is represented by three records, from Inver- 
ness (Aug., 1915), Lincolnshire (Sept., 1916), and Wexford (Sept., 
1914), respectively. 

The male Sowerby's Whale is provided with a pair of large 
triangular teeth in the lower jaw, at about the middle of the 
length of the mouth, on either side. Females have a pair of 
similar teeth in the same position, but, so far as is known, always 
beneath the gum. The three specimens here recorded had their 
teeth concealed by the gum, and were presumably females. The 
Wexford individual was, however, definitely known to be of this 
sex. The Skegness specimen Avas reported to have been 18 feet in 
length, which is unusually long for this species; and it was white 
below — a type of coloration which has been reported by other 
observers, although Sowerby's Whale is often completely black. A 
list containing 33 records of this species has been given by 
Klikenthal (1914, pp. 98, 99). 

The Ziphioid Whales are said to feed principally on Cuttlefish ; 
and it is not impossible that the reduction of their teeth may be 
associated with this diet. Piscivorous animals, such as the 
Delphinid^e and Crocodiles, seem to require a number of sharp 
teeth adapted for holding their prey. In the Ziphioids, teeth of 
this character are obviously not required ; and it may be suggested 
that the absence of teeth is an advantage to them, since the 
suckers of the Cuttlefish would probably attach themselves to 
such convenient pegs, weie these present, and the operation of 
swallowing the prey might thus be rendered more difficult. It 
must not be forgotten, however, that the Sperm Whale, which is 



156 DR. SIDNEY F. HABMER ON CETACEA 

also " teuthophagous," has a specially strong series of teeth in its 
lower jaw. 

(8) Killer or Grampus [Orcinus orca L.). 

A single record, May 1916, from the Sol way Firth. The 
specimen, a male, was unforbunately incomplete, but the part 
remaining was 25 ft. 6 in. long; and the animal must have been 
at least 30 feet long when alive. The species is very common on 
the Norwegian coast (Hjort, 1902, p. 120). 

It has been pointed out by Liitken (1887, p. 367) that a very 
remarkable alteration in the proportions of the fins takes place 
during growth in male Killers. While in young males, and in 
females throughout life, the fins are relatively small and weak, in 
the males all the fins — pectoral, dorsal and caudal — become 
disproportionately lai-ger as the animal grows older. The 
difference between the small pectoral fins of the young male and 
those of the old male is described by this author as being '' per- 
fectly astonishing." These altei^ations in the fins have more 
recently been described and figured by Grieg (1906, p. 9; see 
especially his figs. 2-6, showing alterations in form and size of 
the dorsal, caudal, and pectoral fins). 

The specimen from the Solway Firth was an excellent illus- 
tration of these statements. The dorsal fin was about 5 ft. 6 in. 
in height, while the pectoral limbs were 6 ft. 8 in. long and 
3 ft. 7 in. broad ; thus greatly exceeding in absolute size those of 
a lai'ge Sperm Whale. While the pectoral limbs of young 
animals and of females generally measure about one ninth of 
the total length of the animal, those of the old males ai-e as 
much as one fifth of the entire length. Assuming the correctness 
of this statement of Liitken's, the total length of the old male 
here recorded would have been well over 31 ft. It seems 
probable that the female of this species hardly exceeds half the 
length of the largest males. The failure to recognize the 
occurrence of these changes in the males has i-esulted in the 
introduction of more than one specific name for the Killer. 

Grieg (1906) has given an account of a school of about 47 
Killers which were observed on the Norwegian coast at the 
middle of January, 1904. Four of the females were ascertained to 
be pregnant, and others were accompanied by a calf. The young 
of 2-5 m. in length were regarded as not more than 2 months 
old, and those measuring 3*5 m. as being probably 1 yeai' old. 
Birth and pairing are believed by Grieg to take place during the 
later months, of the year. 

0. area may be easily recognized by its very large teeth, which 
have a basal diameter of as much as \\ or even 1^ in. and are 
10 to 13 in number on each side of each jaw, as well as by its 
strikingly marked black and white (or yellowish) coloration. 
A white patch behind the eye is characteristic, and the white 
area of the ventral surface, although variable in extent, typically 



STRANDED ON THE BRITISH COASTS. 157 

sends up a lateral extension backwards on each side, behind the 
region of the dorsal fin. 

(9) Blackfish or Pilot- Whale {Glohicephcda mehxna Traill). 

Three records in the five years: Hampshire, 1913; Northum- 
berland, 1914; and Cork, 1915; all in March; besides a school 
of about 50 individuals which were stranded at Penzance on 
July 1, 1911. 

The Pilot- Whale reaches a length of at least 26 ft. and is 
usually, completely black. It possesses about 10 large teeth, 
about yV in- in basal diameter, on each side, in the front half of 
each jaw. The "forehead" is enormously swollen, and the 
pectoral limbs are very long and narrow, reaching a length of 
about 4 ft. 6 in. It is well known to associate in large schools, 
which are hunted in the Orkneys, the Faroe Islands, and 
elsewhere. As many as 2000-3000 individuals have been driven 
ashore at one time on the coast of Norway (Hjort, 1902, p. 119) ; 
and a record of 1000 specimens observed at the Lofoten Islands, 
on Sept. 4, 1890, has been given by Grieg (1897, p. 8). 

(10) Risso's Grampus {Grampus griseiis Cuv.). 

Four records : Jersey, Aug , 1913 ; S. Devon, Aug., 1917 ; and 
S. Devon, Nov., 1913 ; the last record consisting of two 
individuals, presumably mother and calf, althougli the sexes 
were not definitely ascertained. It will be observed that all 
these records belong to the S. W. extremity of the British Islands ; 
and there is little doubt that this is a more Southei'U species 
which only just reaches our seas. 

Bisso's Grampus is recognizable by its blunt head, without 
beak; and by the considerable i-eduction of the number of 
its teeth. Of these, none are present in the upper jaw, wiiile the 
lower jaw possesses 4 or 5, of considerable size, with a basal 
diameter of as much as | in., at the front extremity. Adult 
individuals reach a length of about 12 ft. Flower's Memoir 
(1874) may be consulted for information regarding tliis species. 

(11) Bottle-nosed Dolphin [Tursiops truncatus Mont.). 

Ten records: one in February (Scilly Islands, 1915), one in 
May (Merioneth, 1916), and the remainder during the period 
June- August. With the exception of one individual (Essex, 
1914), the determination of which was not quite certain, all were 
from the Southern coast of England, or from the Welsh or 
Lancashire coast. 

The Bottle-nosed Dolphin reaches a length of al)oiit 12 ft., 
and possesses about 25 teeth on each side of each jaw. As in 
some other Dolphins, about two of these, at the front end of the 
series, are very. small and usually remain concealed beneath the 
gum ; while the I'emainder are relatively lai'ge, with a basal 



158 on. SIDNEY F. IIARMER ON CETACEA 

diameter of as much as -^-^ in. As indicateil by its name, the 
head lias a well-marked beak, about 3 in. lonaf. 



(12) White-beaked Dolphin [Lagenorhynchus alhirostris Gray). 

Eight records : one from the North coast of Ireland (March, 
1917), one from the Island of Islay (Oct., 1913), and the remainder 
from the Eastern coast, from Caithness to Lincolnshire — to which 
may be added three specimens stranded in Sufiblk and Kent in 
February, 1918. Although the two species are known to overlap 
in their distribution, it is noteworthy that in this series of 
observations the distribution of the White-beaked Dolphin 
and of the Bottle-nosed Dolphin have not overlapped and are 
complementary to one another. 

Tlie White- beaked Dolphin is characterized, as indicated b}^ its 
popular name, by possessing a well-marked shoi't beak, which is 
white in colour. It reaches a length of rather more than 9 ft. 
Its teeth are about 25 on each side of each jaw, but are distinctly' 
smaller than those of the Bottle-nosed Dolphin, their basal 
diameter being about f'g— fV in. 

(13) White-sided Dolphin {LagenorhyncJms acutus Gray). 

Three records: Fair Island, between the Orkneys and the 
Shetlands (March, 1913); Lincolnshire (May, 1917); Co. Mayo 
(June, 1916). 

This species, which is said to be one of the commonest of the 
Cetacea off the Norwegian coast, where it may occur in schools of 
as many as 1000 individuals (cf. Hjort, 1902, p. 118), may be 
regarded as a boreal species which does not often occur in our 
seas. Most of the specimens previously found have been from 
the Orkneys and other parts of Scotland ; and one or two have 
been noticed fi'om Ireland. So far as I am aware, the Lincoln- 
shire specimen here recorded is the first to have been obtained on 
the English coast. 

The AVhite-sided Dolphin has a characteristic longitudinal 
whitish area on each side, in the middle and posterior half of its 
body. Its teeth are smaller (basal diameter about ^^ in.) and 
more numerous (about 30 to 35 visible during life on each side of 
each jaw) than in the White-beaked Dolphin, from which it 
dift'ers in certain other respects, the following of which may be 
noticed. The beak, which resembles that of L. alhiy^ostris, 
is black. I'he pectoral fins are falcate, with a very convex lower 
boi'der ; those of the other species being blunter and broader, 
and less convex below. These lins, moreover, originate from the 
white part of the body, being connected with the black part of 
the head by a narrow dark streak ; while in L. alhirostris the 
black of the dorsal surface extends as far as the base of the 
flipper (cf. Llitken, 1887, pp. 377, 386, 395). L. aciitus is said 
to reach a length of about 12 feet (Hjoi't, 1902, p. 117). 



STRANDED ON THE BRITISH COASTS. 159 

(14) Common Dolphin {Delphlnus delj^his L.). 

About 20 records, one or two of which liave been not quite 
certain, though probable. Three individuals have been recoi^ded 
frona the Northern part of Ireland (Mayo, Donegal); two from 
the J^orth-E isbei'U coast of Scotland (Inverness, Kincardine) ; 
one, somewhat doubtful, from East Anglia (Suffolk) ; and the 
remainder, either from the English Channel, from Kent to the 
Scilly Islands, or from the entrance to St. George's Channel, on 
both sides, from Cork (1918) to Wexford on the Irish side, and 
on the Welsh coast on the opposite side. None have been 
recorded in the North Sea area, from Forfar to Norfolk ; and 
none on the West coast of Scotland ; which, howevei', has 
provided a curiously small number of records during the whole 
of these observatiotis. The distribution thus indicated is in 
agreement with the supposition that this is an oceanic species 
which is frequently sti'anded on the more exposed parts of the 
coast-lines, but comparatively seldom makes its way into the 
North Sea. The three records from the S. coast of Ireland 
(including one for 1918) were obtained in February, that from 
Inverness in April, and the remainder from August to December. 

The Common Dolphin reaches a length of about 7 ft. 6 in. ; 
aiid is distinguishable by its very long beak and numerous conical 
teeth, of relatively small size, with a bas;il diameter of about 
-f^-^r in. The teeth are more numerous than in any of the otiier 
species here considered, being about 45 in each half of each jaw. 

(15) Common Porpoise {Phoccena phocoina L.). 

Numerous records, indicating, as generally supposed, that this 
is the commonest species in British waters. Of those which were 
certainly determined, the great majority were recorded from the 
East coast of England, and most of them during the period May 
to August. Evidence has been obtained in support of the belief 
that the Common Porpoise gives birth to its young in the early 
summer, and that the length at birth is fi'om 2 ft. to 2 ft. 6 in. 
According to Prof. Meek (1918, p. 197), the occurrence of 
Porpoises near the coast during July and August may be 
reo'arded as an inshore migration for parturition and pairing. 

This species differs from all others found on the Bi-itish coasts 
in the form of its teeth, which instead of being conical, as in the 
majority of the species, are broadened at the free end ('-spade- 
shaped"), although the exact form of the broadened part 
is variable. The teeth undergo a considerable amount of 
thickening as growth proceeds, and the broadened blade is not 
infrequently worn away in some of the teeth. About 25, 
or rather more, are present on each side of each jaw ; though 
usually two, at the anterior end of the series, remain small a,nd 
are concealed below the gum. The length of the adult is about 
5 ft. 6 in. — and this is distinctly the smallest of the British 
Cetacea. The head is not provided with a beak. 



160 dr. sidney f. harmer on cetacea 

Memoirs cited, 

Allen, G. M., 1916.— "The Whalebone Whales of New England." 

Mem. Boston Soc. Nat. Hist. viii. No. 2, p. 105. 
Andrews, R. 0., 1916.— '' Monographs of the Pacific Oetacea," II. 
'• The Sei Whale {Balcenoptera horealis Lesson)." Mem. 
Amer. Mus. Nat. Hist. (n. s.) i. part vi. p. 289. 
BuRFiELD, S. T., 1913.— "Belmullet Whaling Station." Rep. 

82nd Meeting (Dundee, 1912) Brit. Assoc, p. 145. 
CocKS,_A. H., 1884.— "The Finwhale Fishery on the Coast of 

Finmark." The Zoologist, (3) viii. p. 455. 
Cooks, A. H., 1887.— "The Finwhale Fishery of 1886 on the 

Lapland Coast " The Zoologist, (3) xi. p. 207. 
CoLLETT, R.., 1906.—" Nogle Meddelelsen om Naebhvalen 

(Hyperoodon), og Hvidfisken {Delphinapterus).'' Bergens 

Mus. Aarbog, 1906, No. 6. 
CoLLETT, R., 1912.— "Norges Pattedyr." Kristiania, pp. 543-. 
Flower, [Sir] W. H., 1874.— "On Risso's Dolphin, Grampus 

griseus (Cuv.)."' Trans. Zool. Soc. viii. p. 1. 
Grieg, J. A., 1897. — " Nogle cetologiske notiser." Bergens Mus. 

Aarbog, 1897, No. 6. 
Grieg, J. A., 1906. — "Nogle uotiser fra et spfekhuggerstseng 

ved Bild^str^mmen i januar 1904." Bei-gens Mus. Aarbog\ 

1906, No. 2 
Hamilton, J. E., 1915.— " Belmullet Whaling Station." Rep. 

84th Meeting (Australia, 1914) Brit. Assoc, p. 125. 
Hamilton, J. E., 1916.— " Belmullet AYhaliug Station."' Rep. 

85th Meeting (Manchester, 1915) Brit. Assoc, p. 124. 
Harmer, S. 1''.—" Report[s] on Cetacea stranded on the British 

Coasts during [1913-1917]." Published by the Trustees of 

the British Museum. 

1914. No. 1, Cetacea stranded during 1913. 



1915. 


No. 2, 


?? 


?J 


?) 


1914. 


1916 


No. 3, 


?5 


5« 


59 


1915. 


1917. 


No. 4, 


55 


9> " 


?) 


1916. 


1918. 


No. 5, 


5? 


?; 


9? 


1917. 


Harmer, S. F. 


, 1915.— 


"On 


8pecimens 


of 


Cuvier's 



Whale 

{Ziphius cavirostris) fi^om the Irish Coast.'' Proc. Zool. 

Soc. 1915, p. 559. 
Hjort, J., 1902. — " Fiskeri og Hvalfangst i det Nordlige Norge." 

Aarsberetn. vedk. Norges Fiskerier for 1902, i Hefte, Bergen 

(udgivet nf Norges Fiskeristyrelse). 
Kukenthal, VV., 1914 — " Untersuchungen an Walen (Zweiter 

Teil)," VI., " Zur Kenntnis des Mesoplodon hidens (Sow.)." 

Jen. Zeitschr. li. p. 93. 
LiLLiE, D. G., 1910. — "Observations on the Anatomy and 

General Biology of some Members of the Larger Cetacea." 

Proc. Zool. Soc. 1910, p. 769. 
LiJTKEN, C. F., 1887. — " Kritiske Studier over nogle Tandhvaler 

af Slaegterne Tiirsiops, Orca og Lagenorhynchus ." Vid. 



STRANBED ON THE BRITISH COASTS. 161 

Selsk, Skr. ((5), naturvid. og math. Aid., Bd. iv. No. 6 

(Copenhagen), p. 337. 
MeeKi, a., 1918. — "The Reproductive Organs of Cetacea." Journ. 

of Anat. Hi. p. 18G. 
MiLLAis, J. G., 1906. — "The Mammals of Great Britain and 

Ireland," vol. iii., London (Longmans, Green & Co.). 
Munsterhjelm, L., 1915, — " Anteckningar om Hyperoodon 

rosir«fi<s(Miill.)gjorda under en ishavsi'esa sommaren 1910." 

TromS(j!> Museums Aarshefter, xxxvii. 1914, p. 1. 
Murray, [Sir] J., and Hjort, J., 1912.— "The Depths of the 

Ocean." London (Macmillan & Co.). 
Owen, [Sir] R., 1840-1845.— "Odontography," 
Sars, G. O., 1881. — "Fortsatte Bidrag til Kundskaben om vore 

Bardehvaler," Forh. Yid. -Selsk. Christiania, Aar 1880, 

No. 12. 
Southwell, T., 1883. — "On the Beaked or Bottle-nose Wha^B 

( Hyperoodon rostratas). Trans. Norfolk and Norwich Nat. 

Soc. iii. p. 476. 
Thompson, D'A. W., 1912.— "Whales, Seals and Sea-Serpents," 

in " Science of the Sea." Edited for the Challenger Society 

by G. H. Powler, ch. xii, pp. 383-402, London (John 

Murray). 
True, F. W., 1904.— "The Whalebone Whales of the Western 

North Atlantic compared with those occurring in European 

Waters, with some Observations on the Species of the North 

Pacific." Smithson. Contr. xxxiii. p. 1. 
True, F. W.; 1910.— "An Account of the Beaked Whales of the 

Family Ziphiidse in the Collection of the United States 

National Museum, with Remarks on some Specimens in 

other American Museums." U.S. Nat. Mus., Bull. 73. 
Turner, [Sir] V\^., 1871.— "On the Capture of a Sperm Whale 

on the Coast of Argyleshire, with a Notice of other Specimens 

caught on the Coast of Scotland." Proc. Roy. Soc. Edinb. 

vii. p. 365. 
Turner, [Sir] W., 1872. — "Additional Notes on the Occurrence 

of the Sperm-Whale in the Scottish Seas." Proc. Roy. Soc. 

Edinb. vii. p. 632. 
Turner, [Sir] W., 1886.— "On the Occurrence of the Bottle- 
Nosed or Beaked Whale {Hyperoodon rostratus) in the 

Scottish Seas, with Observations on its External Characters." 

Proc. R. Phys. Soc. Edinb. ix. 1885-1888, p. 25. 
Turner, [Sir] W., 1904.— "The Occurrence of the Sperm Whale 

or Cachalot in the Shetland Seas." Ann. Scott. Nat, Hist. 

1904, p. 4. 
Turner, [Sir] W., 1912.— "The Marine Mammals in the 

Anatomical Museum of the University of Edinburgh." 

London (Macmillan <fe Co.). 
Van Deinse, A. B., 1918, — "Over de Potvisschen in Nederland 

gestrand tusschen de Jaren 1531-1788," Zool. Meded. R. 

Mus. Nat. Hist. Leiden, Deel iv., Afl. 1, p. 22. 
Prog. Zool. Soc— 1918, No. XI. 11 



ON THE VARIATION OF THE PIT-VIPER. 163 



8. On the Variation of the Pit- Viper, Lacliesis atrox. 
By Miss Joan B. Procter, F.Z.S. 

[Received March 19, 1918 : Read April 9, 1918.] 

(Text-figures 1-5.) 

Index. 

Page 
Variation and ^Etiology (Evolution, heredity, etc.). Variation 

in Lacliesis atrox; derivation of markings 167-180 

Systematic (New names or changed names, or Varieties of 
Xi. atrox ; revisions, discussions, or elucidations affecting 
any part of the system, to be indexed under the names)... 163-167, 180 

The distinction of the forms, whether they he regarded as 
species or as varieties, which cluster around the tropical American 
Pit-'Viper, Lachesis atrox L., and of which the principal are 
L. lanceolatus Lacep^de, L. jararaca Wied, and L. jararacussu 
. Lacerda, is a subject which needs renewed investigation, especially 
in view of Dr. Vital Brazil's recent publications*, in which he 
claims specific rank not only for X. lanceolatus but also for 
L. jararacussu, which has generally been regarded as a mere 
colour variety. 

Mr, G. A. Boulenger, who has kindly helped me with advice in 
my study of reptiles, suggested to me that I should take up this 
investigation, and he has given me not only access to the collection 
in the British Museum, of which he is in charge, but the benefit 
of his experience, especially as regards the bibliography of the 
subject. My best thanks are due to him for these favours, 

I. Historical. 

The nvxmber of names which Mr. Boulenger has quoted under 
the synonymies of L. atrox and L. lanceolatus is very great, but 
for my present purpose it will be sufficient to discuss only the 
principal, such as have had more general currency, or which are 
accompanied by figures enabling me to form an opinion on the 
forms for which they ai-e intended. 

The earliest name is that of Coluber atrox Linnfeus, Mus. Ad. 
Frid. (1 754) pi. xii. fig. 2. The specimens are described as having — 
ventrals 200 and 196, subcaudals 70 and 67 pairs, and scales with 
'■•carina elevata." However, Mr. L. G. Andersson, Oat. Lin. type- 
spec. Sn. (Bih. Sv. Vet.-Ak, Handl. xxiv. iv. No. 6) p. 19, has 
been abFe to supplement this definition, as he has had access to 
the type-specimens in the Stockholm Museum, and he finds that 
the keels on the scales " are low and extend nearly to the tip 
of the scales," thus agreeing with Mr. Boulenger's L. lanceolatus, 

* La Defense coutre I'Ophidisme. Sao Paulo, 1911, 8vo. 2nd edition, 1914. 

11* 



164 MISS JOAN B. PROCTER ON THE . 

not with liis L. atrox. He counts 204 and 200 ventrals (v.), 
67 and 70 subcandals (c). Both descriptions appear to corre- 
spond with two specimens in the British Museum collection, from 
Martinique (Cat. Shakes, iii. p. 536, spec. ?■ & s). 

Goluher ^ffl?iceoZ«itts Lacepede, Hist, des Serp. (1789) p. 121, is 
based on specimens -in the Paris Museum: 228, 225 v., 61, 59 c. 
Habitat Martinique and perhaps Dominica and Cayenne. Figure 
worthless, probably from a bleached specimen. Dumeril and 
Bibron (Erp. Gen. vii. 1854, p. 1505), under Botlirops lanceolatus, 
do noC give us any particulars concerning the type -specimen, 
which -they must have examined, but the fact that the " Vip6re 
jaune de la Martinique," also found in St. Lucia and Dominica, is 
the form intended. There can be no question as to tlie appli- 
cation, of the name Imiceolatns, the Fer-de-lance, wdiich must be 
regarded as a strict synonym of L. atrox of Linnaeus according to 
Andersson. ' 

Cophias jararaca Wied, Abbild. Nat. Bras. (1825), from 
E. Brazil, 193-201 v., 59-68 c. The description and figure 
indicate a snake similar to L. atrox, but with markings consisting 
of dark brown, darker-edged transverse bands, narrower on the 
back than on the sides, such as is figured by Jan, in Icon. Oph. 
47me livr. pi. lii., and corresponding to several examples in the 
British Museum collection from Rio Janeiro and Para (Cat. Sn. 
iii. p. 537, spec. ?', p. 539, sj)ec. v). This variety must be the 
prevalent form in the Province of Bahia, as the following 
descriptions and figures of Wagler in Spix's Serp. Bras. (1824) 
are evidently referable to it. Bothrops megcera, p. 5,0, pi. xix. : 
195 v., 53 c. This figure shows dark olive upper parts with 
dark, darker-edged transverse bands and immaculate ventrals. 
B. fioria, p. 52, pi. xx. : 201 v., 65 c. A uniformly coloured 
specimen which may be taken to have lost its markings. 
B. leucostigma, p. 53, pi. xxi. fig. 1 : ? v., 66 c. Brownish, with 
darker transverse bands, ventrals powdered with grey. B. tessel- 
latus, p. 54, pi. xxi. fig. 2: 190 v., ? c. Dorsal markings as in 
B. leucostigma, ventrals checkered with brown. B. tamiatus, 
p. 55, pi. xxi. fig. 3. Colours lighter than the above, transvei'se 
bands always double. 

Bothrops jararacibssu Lacerda, Leg. sur le ven . des Serp. ( 1 884) p. 9. 
From the province of Rio de Janeiro. An adequate description 
of the striking markings of the Jararacussu is given (black above 
with yellow markings, longitudinal on the head, and oblicjuely 
pronged forks on the sides, yellow beneath spotted with black). 
Similar to the B. atrox figured in Jan, Icon. Oph, (47me livr. 
pi. ii. fig. 3), and to two specimens in the British Museum 
collection from Sao Paulo, presented by Dr. Brazil. 

Having thus indicated the sources to which I have referred in 
order to fix the exact meaning of the principal names which have 
had currency in the past, I will review the various opinions which 
have been expressed in the principal works on Ophidia. . 



VARIATION OF THE PIT- VIPER. 165 

Schlegel (Phys. sei-p. (1837) pp. 532, 535, 537) recoginses three 
species: — 1. Trigonocephalus jararaca, from < Sao Paulo, Sta. 
Oatharina I., Sta. Cruz de la Sierra: 172-204 v., 44-62 c. From 
the description, this snake is the true Jaiaraca of Wied. 
2. T. atrox: 190, 196 v., 60, 68 c. Scales surmounted by a strong- 
keel with tubercular tendency, in which respect it does not agree 
with the Linnean type specimen. Habitat N. Brazil, Dutch and 
French Guianas. 3. T. lanceolatus : 271, 220 v., 60, 60 c. Mar- 
tinique and St. Lucia. Scales surmounted by a simple keel. 
Colour of ventrals fi clear greenish yellow. The description 
corresponds to the type-specimen of L. atrox as described by 
Andersson. Schlegel maintains that T. lanceolatus has a higher 
number of ventrals than the two preceding, but the number 271 
is probably due to some mistake. 

Dumeril and Bibron (Erp. Gen. (1854) pp. 1505, 1507, 1509) 
describe three species :■ — Bothrops atrox^B. lanceolatus, B.jararaca. 
The only diiferentiatioh made between the first two in the 
" Tableau Synoptique des Especes du Genre Bothrops^' (p. 1504), is 
that the venti-als of B. atrox are spotted, and those of B. lanceo- 
latus unicoloured. In describing B. atrox, liowever, they mention 
that the scales are more strongly keeled. B. jararaca is flistin- 
guished from the preceding by a relatively obtuse canthus rostralis 
and larger scales on the snout. The description corresponds with 
that of the Jararaca of Wied. 

B. atrox. 29-32 scales. Habitat Dutch and French Guianas. 
B. lanceolatus. 271, 240, 220 v., 68, 64, 60 c.; from Marti- 
nique, Dominica, St. Lucia. 

B.jararaca. 172, 195, 204 v., 44, 62, 65 c. : from Sao Paulo, 
Sta. Catharina I., never in Surinam. 

The number of venti-als given by Dumeril and Bibron do 
not seem very reliable, and 271 for B. lanceolatus is probably 
copied from Schlegel's statement, which there is good reason to 
question. 

Jan, Icon. Oph. 47me livr. PI. i. : Bothrops lanceolatus (Merr.). 
Antilles? Geneva Museum. The specimen figured has 29 rows 
of scales. Markings irregular, but similar to those of two spe- 
cimens in the British Museum collection from Martinique (Cat. 
Sn. p. 536, spec, r, s). — PI. ii. : B. atrox L., var. dirus Jan, from 
Buenos Ayres. Turin Museum. 25-27 rows of scales, with keels 
similar to those figured for B. lanceolatus. The markings repre- 
sented in fig. 3 nnn. are matched by two specimens in the 
British Museum collection, from Sao Paulo (presented by 
Dr. Brazil), and, as before mentioned, agreeing with those of the 
Jararacussu of Lacerda. — PI. ii. fig. 4 : var. tesselatus Neuw. 
Milan Museum. Another form of dorsal markings, similar to a 
specimen in the British Museum collection, from St. Lucia (Cat. 
Sn. p. 536, spec. v). — PI. iii. : B. jararaca Neuw. Freyburg 
Museum. An excellent figure of the typical Jararaca ; 25 rows 
of scales. 



166 MISS JOA^^ B. PROCTER ON THE 

A. E. Brown, in a paper in the Proc. Acad. Philad. 1893 
(pp. 435-6), describes three species, but adds that he doubts their 
right to more than a varietal distinction : — 1. Bothroj^s atrox, 
described from two specimens from British Guiana; 196, 195 v., 
42, 59 c, 27, 27 rows of scales, which are broad, with a high 
short keel not extending to the tip. Abdomen unspotted. 
2. B. lanceolaius, described from two specimens from Martinique; 
199, 200 v., 69, 69 c, 25, 25 sc. Scales longer than in B. atrox, 
and bearing a simple keel to the tip. 3, B.jararaca, described 
from one specimen from. Brazil ; 206 v., 57 c, 27 sc. ; besides the 
annulated markings, Mr. Brown notes that the canthus rostralis 
is less sharp and the scales on the snout are larger than in the 
preceding species, a fact which was also observed by Dumeril 
and Bibron. 

G. A. Boulenger, Cat. Sn. vol. iii. 1896, pp. 535-537. 

Lachesis atrox. 161-216 v., 47-73 c. 

L. lanceolatus. 180-240 v., 46-70 e, 

Mr. Boulenger finds that the only distinctive character between 
these species is that of scale-structure. In the former the scales 
are "strongly keeled, the keels on the posterior part of the back 
very high, swollen in the middle, and much shorter than the 
scale." In the latter the scales are merely " sharply keeled," the 
keel extending nearly to their extremity. He also describes the 
ventrals of L. lanceolatus as yellowish, uniform, or powdered, or 
spotted with brown, and in this respect many of the specimens 
in the British Museum collection certainly disprove the state- 
ment made by other authors, that the ventrals of this snake are 
constantly immaculate. He adds that these species may have to 
be united, as some specimens of L. lanceolatus "approach L. atrox 
in the swelling of the scales at the base of the keels, and are thus 
intermediate between the two species." Cophias jararaca and 
Bothrops jararacussu ai'e placed in the synonymy of L. lanceo- 
latus. 

V. Brazil, Def. contre I'Ophidisme (2nd ed, 1914), pp. 78, 84, 88. 

L. lanceolatus. 195-200 v., 50-53 c. Tropical America. 

L. atrox. 202 v., 55 c. Tropical America, less abundant than 
the above in the southern states of Brazil. 

L. jararacugu. 170-176 v. Brazil (Sao Paulo and Rio). 

In describing the first two species Dr. Brazil maintains that 
one of the main points of difierence between these snakes lies in 
the " systeme de coloration " of L. atrox, of Avhich he says : " La 
coloration du fond, sur lequel se dessinent des figures pareilles a 
celles eonstatees dans I'espfece prc'cedente {L. lanceolatus) est d'un 
gris rougeatre, parfois un ton gris cendre. Cette comhinaison de 
coulears donne un aspect veloute a Vanimal, ce qui permet de le 
reconnoitre a premiere mte." The ground-coloui> of L. lanceolatus 
is described as " vert tres fonce, gris et quelquefois jaun^tre," 
and the abdomen "vert fonce, parseme de taches jaunes," whilst 



VARIATION OF THE PIT-VIPER» 167 

that of L. atrox is of a " tr^s beau jaune clair marquete sur les 
bords de noir ou de gris fonce"*. 

In examining a large number of these snakes I have found the 
ground-colour to be equally variable in both ; the ventrals may 
be yellow, yellow checkered with black, or of that greenish colour 
produced by a powderiiig of black over yellow, in either L. atrox 
or L. lanceolatus. It was ther-ef ore not quite clear to me why 
Dr. Brazil considers the general aspect and coloration such striking 
points, or how the latter should produce a velvety appearance in 
L. atrox. Mr. Boulenger has looked into this matter with me, 
using specimens of L. atrox and L. lanceolatus named and sent to 
him by Dr. Brazil himself. There certainly is a difference in the 
appearance of these specimens, but this is not due in any way 
to the markings, which are strikingly similar. The specimen 
marked " at7'ox" has, however, the very prominently keeled scales 
mentioned before, and Mr. Boulenger has pointed out to me that 
the " velvety " appearance of this snake is entirely due to the 
consequently greater breaking up of light upon its dorsal surface. 
Dr. Brazil's L. atrox is therefore the same as Mr. Boulenger's. 

L.jararacussu. — Dr. Brazil maintains the specific rank assigned 
to the Jararacussu by Lacerda : first, on account of its striking 
black and yellow markings ; secondly, because of the comparatively 
low number of its ventrals ; and, thirdly, because of its much 
more triangular head. With regard to the first point, there are 
specimens in the British Museum collection which can be graded 
to form a transition series from the typical L. atrox to the 
Jararacussu, a specimen from W. Ecuador being exactly inter- 
mediate between the two types of markings. With regard to the 
second point, the ventrals of the Jararacussu are as a rule fewer 
than those of L. atrox. Dr. Brazil mentions 170-176, but two 
specimens received from him have 180 and 184. 180-240 ventrals 
is, however, the range given for L. lanceolatus in Mr. Boulenger's 
Catalogue of Snakes, and, as I have failed to detect any structural 
differences in the shape of the head or in the scaling, I think 
that the Jararacussu cannot be regarded as more than a colour 
variety of L. atrox L. 

II. FORBI AND LePIDOSIS. 

As I think I have shown in the annexed table that there is no 
correspondence between variations in markings and the number of 
ventrals and caudals, I will describe the differences in form and 
lepidosis which have led to the distinction of species. 

The most important is that of the two types of scaling found 
in L. atrox. It has been generally accepted that the high short 

* The coloured plates which accompany Dr. V. Brazil's descriptions do not convey 
these supposed difierences, and it is well to point out that but little care has been 
bestowed on the rendering of the markings. I think a L. lanceolatus with only 
about 10 markings on the body, as figured, to be an impossibility. 



168 MISS JOAN B. PROCTER ON THE 

keel is pi-op.er to L. atrov and the low long keel to L. lanceolatus. 
Dr. Anrlersson has shown, however, that the latter form is really 
the Coluber atrox of Linvijeus. so that if the former is maintained 
as a distinct species on account of its scale-structure, it will be 
necessaiy to alter the name which has usually been given to it. 
I suggest that of L. ajjinis Gray, as the specimens of Bothrops 
affinis (Catalogue oi Snakes (1849), p. 7, specimens / & o), which I 
have examined, answer the definition, and this appears to be the 
earliest name which can be applied to it. 

The dorsal scale of the typical L. atrox L. is of a long narrow 
diamond shape, usually about twice as long as broad, and bearing 
a simple keel extending to the apical pits in the tip of the scale. 
That of L. affinis is broader in proportion, more rounded, and 
sometimes truncate behind ; it bears a short keel, supported upon 
an extremely convex area, leaving only a naiTOw margin of flat 
scale. This convexity of the scales is so pronounced in some speci- 
mens that they can be distinguished from the jJreceding form by 
touch alone. The scales of other specimens, however, ai-e inter- 
mediate in type ; they may be of the narrow form with the long 
keel distinctly swollen at the base, or, while maintaining the long 
keel of atrox, they may be both broad and convex as in affinis. 
Other individuals present scales of both the extreme types. On 
examining the middle third of the body in one of these, I find that 
the median dorsal scales are of the high short type, and the lateral 
dorsals of the long low type. Further examination shows that, 
in every case, the anterior part of the snake is of the atrox 
type and the posterior of the affinis type. The evolution of the 
scale-structure is thus clearly shown. The long low-keeled scale 
gradually broadens, whilst its keel swells along its base and 
shortens, until it becomes completely transformed into the affinis 
type. This transformation is, as described above, completed upon 
the median dorsal scales sooner than upon the laterals. 

As regards the snout of L. jararaca, its shape is very slightly 
diflferent from that of L. atrox, in that the canthus lostralis is 
somewhat more obtuse, and the scales of the upper surface are 
slightly larger than those on the vertex. 

These characters are not very distinct, as several specimens of 
the typical L. atrox pi'esent snouts of a similar form ; in fact, 
one specimen (Cat. Sn. iii. p. 536, spec, v) is indistinguishable 
from L. jararaca (spec, g, Cat. Col. Sn. [1858] p. 226) in this 
respect. Dumeril & Bibron, and A. E. Brown, both mention 
this point as one of the distinguishing features between the 
two forms ; and Schlegel (Phys. Serjj.), in his figure (pi. xix. 
fig. 1) of the dorsal view of the head of the Jararaca, represents 
the difference between the scales on the snout and those on the 
rest of tlie head as very considei'able. This is certainly a point 
of variation which I cannot consider proper to the Jararaca 
alone, having found no fewer than six young specimens of 
L. atrox with this chai^acteristic. 

The different types of form and lepidosis described above are : 



VARIATION OF THE PIT-VIPER. 'lf)9 

therefore so incon^staut, and so com^jletely connecterl by inter- 
mediate forms, that they cannot in themselves be considered 
sufficient for the distinction of species. 

III. System of Markings. 

Before discussing the evolution of markings and their individual 
varia,tion, I will outline the general system of arrangement of 
markings common to all colour varieties of L. atrox. As a working 
hypothesis on this vsubject, Mr. Boulenger drew my attention to 
Dr. Zenneck's paper " Die Zeichnung der Boiden "*, and I have 
found that his theoi-y — that the dorsal and lateral markings of 
Boids are made up of four paired longitudinal series t — is equally 
applicalde in the case of L. atrox. Of these, only thi-ee are 
usually present, but in some specimens the fourth is present upon 
the head. It is the constant relationship between the spots 
of the series of one side of the snake which forms the regular 
pattern, but those of one side are quite independent of those 
on the othei', thus accounting for the asymmetrical appearance 
of the markings on the median dorsal line, which is sometimes 
very marked J. 

When present on the head, the first or dorsal series commences 
as a streak, or a few broken spots. Occasionally it forms a 
A-shaped marking with its fellow. On the body this series is 
usually the broadest and most marked, and, in individuals 
where it tends to disappear, it is always the last to go. The 
dorso-lateral or second series, when present on the head, may 
either consist of an oblique narrow streak or of broken spots of 
this streak, originating above the ocular shield, or of a cross-bar or 
blotch in front of the eyes. This second series is rather unstable 
on the V>ody, for though it may be of equal development to the 
dorsal, it is moi-e often hardly discernible and certain spots 
frequently become confluent with their neighbours of the first 
series. In some forms the markings of these two paired series 
form a single dark triangular area on the head, but are more 
often irregular and indistinct. 

The lateral series is always present on the head of those snakes 
exhibiting dorsal markings. It forms a bioad black streak from 
the posterior boixler of the eye to the commissure of the jaws, 
usually passing through the sixth and seventh labials. Rays 
proceeding from it may be present on the other upper labials, or 
may take the form of isolated spots. The post-ocular streak is 
always separated from the superciliary streak by a narrow ai'ea 
of the lighter ground-colour, and is sometimes also outlined in 



* Tiibing eu Zool. Arb. iii. 

t These four series are termed Dorsal (D.), Dorso-lateral (D.L.), Lateral (L.), and 
Veiitro-lateral (V.L.) by Mr. Boulenger, 'Snakes of Europe' (1913), p. 30. This 
lettering is followed in mj' text-fig. 1. 

X Mr. Boulenger has drawn attention to this bi-lateral asymmetry in his op. cit. 
p. 33, and he mentions a specimen of Lachesis alternatus with 24 markings on the 
left side and 27 on the right. 



170 



MISS JOAN B. PROCTER ON THE 



cream -colour, which brings it out very sharpljc On the body this 
series is well defined. 

The fourth or latero-ventral series is never present]on the body, 
and not always upon the head. It is evidently a most primitive 
character, vvhieli has disappeared in many specimens. In the 

Text-figure 1. 








-O-L 




tiV 



./KmBBm_ 



.♦....-..#. 



D-L-- 
— L---- 



most marked form this series only consists of three or four blapk 
spots on the lower labials, and one larger spot which has become 
confluent with the post-ocular streak. This latter spot is the 
last of the series to disappear, and is entirely the cause of the 
apparent broadening and lengthening of the post-ocular streak in 



VARIATION OF THE PIT-VIPKR. 



171 



many snakes. This series must not be confounded with the outer 
ventral series, which commences on the chin-shields and is usually 
distinctly marked throughout the body. 

The markings develop upon regular lines, and the degree of 
this development varies on different regions of the body. That 
on the anterior end of the body is the most primitive, as I shall 
endeavour to show later on, but markings immediately behind 
the head are always irregular. 

In Lachesis atrox the spots of the series on each side of the 
body form themselves into distinct alteinating groups, which I 
shall call A and B. Group B is less stable than A, and frequently 
disappears altogether, either tempoiarily in the evolution of the 
markings or permanently. 

Text-figure 2. 
Group A. Group B. 

I \ 







♦ 



On one snake the relative development of groups A and B is 
not always constant, but it is always alike on both sides of 
the body. 

Since group A is usually the best developed, it is this group 
which I have counted in giving the numbers of markings in the 
table of statistics. This number is not the same for the two sides 
of many snakes — another reason for the faulted appeai'ance in 
the general dorsal pattern. 

The dorsal markings of L. atrox are therefore made up on each 
side of three longitudinal series of spots, which in turn form 
themselves into alternating groups A and B. Each of these 
consists of six primitive spots, which, according to their presence, 
absence, or confluence, determine the variety of marking pro- 
duced. In oi'der to follow their evolution and variation, I have 
numbered those of each group in order of their stability (see 
text- fig. 2), so that each may be referred to by a definite 
formula. 

The ground-colour appears to be formed by the varying pro- 
portions of a black (or dark brown) and a yellow pigment, the 
black being the more superficial. When the shade is gi-eyish, 
there is a smaller proportion of yellow pigment. The usual brown 
tints are composed of a very fine powdering of the dark pigment 
over the yellow. Quite apart from the regular markings, the scales 



172 MISS JOAN B. PROCTEK ON THE 

often each show a delicate arrangement of black pigment standing 
out from the ground-colour and having the same pattern effect 
as the fine dendritic marks seen in many rocks. This arrange- 
ment is often exactly repeated from one scale to another. When 
this style of markings is pi-esent on the clear yellow of the 
ventral shields, it produces a dark green effect, merging, when 
seen at a distance, into the ground-colour. 



IV. Individual Variation and Evolution of Markings. 

Individual Variation. 

As a starting-point in the study of the markings and colour 
varieties of L. atrox, it was suggested to me by Mr. Boulenger 
that I should make an examination of a female and her 26 young 
from Trinidad (received from Mr. Urich) in order to ascer-tain 
the amount of individual variation among specimens unqviestion- 
ably pertaining to one form only. 

The young snakes range from 160 to 300 mm. in length, and 
are more brightly marked than their mother. Their ground- 
colour is a soft shade of brown, except the tip of the tail, which 
is a pale yellowish. Particulars as to the number of ventrals 
and subcaudals'etc. will be found in the annexed table. In the 
majority of cases, the pattern on each side is made up in the 
following way, as shown on text-fig. 3 : — 

Group A consists of the six primitive spots arr-a"nged in a 
pyramid (see text-fig. 2, p. 171). A large triangular or rather 
trapezoid spot (A 1, dorsal series) forms the apex, and rests 
upon two small rhombic spots (A 5, 6, dorso-lateral series), which 
are sometimes confluent with it. These rest upon three similar 
spots, the outer ones being somewhat the larger (A 2, 4, 3, lateral 
series). This triangular marking is margined in cream-colour, 
and encloses an area darker than the surrounding ground-colour, 
a state of things which seems to suggest a centripetal aggregation 
of pigment. Group A, though equal in size to group B, is in 
every case darker and more marked. 

Gi'oup B is also triangular in form, but the base coincides with 
the mid-dorsal line, instead of the apex as in group A. The said 
base is composed of three oval spots (B 5, 1, 6, dorsal series), the 
middle one being much the largest. Three more small oval spots 
enter the remainder of this triangle, the largest at the apex 
(B 3, 4, dorsolateral, B 2, lateral). (Text-fig. 3, A.) 

When the markings of the two sides of the snake correspond 
exactly, the general dorsal design is a single string of light- 
spotted ovals on a dark ground ; this is formed by the paired 
groups B having their bases applied together. They appear oval 
rather than diamond-shape, on account of the curved border of 
the ground-colour of these groups. If the markings should alter- 
nate with each other, a light wavy band on a dark ground is the 
general result. Usually, however, a single specimen exhibits 



VARIATION OF THE PIT-VIPER. 



173! 



iriany different combinations of these forms, owing to the different, 
number of markings on tlie two sides. In most specimens mai-k- 
ings of one or more series are present on the head, and in several 
the latero- ventral series is very marked. 

Text-fiffui^e 3. 



















Many specimens do not possess all the primitive spots described, 
especially upon the posterior pni-t of the body, where the pattern 
becomes modified. In the first stage of this modification A 5 
and 6 become confluent with A 1, which tends to divide into two 
triangular spots ; the spots of group B, with the exception of 
B 1 and 2, merge into the background (text-fig. 3, c). Some 
specimens are of the more primitive type anteriorly only, whilst 
others have the markings of the greater part of the body of the 
slightly modified type, and become, still further altered -on their 



174 MISS JOAN B. PROCTER ON THE 

posterior parts. One specimen, however (see Table, Trinidad 1 A), 
is inclined towards the Jararaca type of marking. Anteriorly 
A 1 is divided, each spot being confluent with A 5 and 6 ; A 4 is 
absent, and of group B only an occasional- B 1 is present (text- 
fig. 3, d). At about tlie middle of the snake the twin spots 
A 1, 1, have become some distance apart, and are in sufficiently 
close proximity to A 2 and 3 to give the appeai-anee at a distance 
of paii'ed transverse bands, or, since the intermediate space is 
darker than the surrounding ground-colour, of a single dark- 
edged band ; these bands, which may pair with those of the 
opposite side or not, are very little more than their own width 
apart. 

In all specimens the pattern tends to break up into a spotted 
type just before the tail, all the spots being more or less equal in 
size and distribution. It consists of A 1, 1, 4, 2, 3, and B 1, 
3, 4, and 2. 

In the markings of the mother, A 1, 2, 3, 5, and 6 are con- 
fluent, forming a somewhat truncate chevron, in the middle of 
which A 4 is sometimes situated. Group B is made up of the 
six primitive spots, all except B 1, 2, however, being indistinct 
(text-fig. 3, b). On the whole, the individual variation shown by 
these 27 specimens is very slight : the most apparent differences 
between them are due not to variation of the actual markings, 
but to the inconstant I'elationship of those of the two sides of 
the snake. 

1 have since examined another mother and young, from Anda- 
goya, Colombia (received from Dr. Spurrell), and in this case 
there is no variation at all among the young. 

The markings of both families are of the most primitive type 
occurring in Z. atrox^ all other forms being evolved from it, as 
will be shown presently. 

Evolution^ 

In the course of evolution, these primitive markings become 
modified in the following way, which is the same for all forms of 
L. atrox. 

A 1 tends to divide transversely to the axis of the body, and 
A 5 and 6 become confluent with it. B 5 and 6 merge into the 
wround-colour, A 4 and B 3 and 4 disappear (this stage has 
already been described in the variations of the young snakes), 
A 1 then divides completely, but is still distinct fi-om A 2 and 3._ 
A 4 disappears, and the whole of group B tends to merge into 
the background. The spots A 1, 1 then become a considerable 
distance apart, narrow, and in- close proximity with A 2 and 3, 
which, since the interspace is darker than in group B, thus 
produce the effect of a dark, darker-bordered, transverse band. 
Except for an occasional B 1, this group is pale and quite 
indistinct. At this point the bands, which may or may not pair 
with those of the opposite side, are slightly more than their own 



VARIATION OF THE PIT-VIPER. 



175 



width apart. A 1, 1 and A 2, 3 now become confluent, forming 
the dark unbroken marginal lines of the transverse band which 
constitutes group A (text-fig. 4, a). These lines may be nearer 
together on the back than on the sides, and are sometimes much 
serrated. The bands, i. e. groups A, gradually become further 
apart from each other, with the result that the markings are 
slightly fewer in number than in the primitive type. For the 
present I shall call this annulated form of marking Type II. It 
constitutes a distinct branch in the direct evolution of markings 
(text-fig. 3, a). Type III. evolves directly from the primitive 

Text-figure 4. 






■|^|i^^M^M 






arrangement, in exactly the same way as Type II. up to the point 
whei^e A 1 is completely divided, but distinct from A 2 and 3. 
A 1, 1 are, however, squarish, and form a square group with A 2 
and 3, which they equal in size. In the course of evolution A 4, 
B 2, 3, and 4 reappear (text-fig. 4, b), and are also all of equal 
size. In this type A 4 belongs to the dorso-lateral series, being 
situated immediately above its position in the lateral series in 
Type I. In the evolution from Type I. to Type III. the series 
equalise, the dorsal narrowing and the dorso-lateral broadening. 
The general pattern is formed, therefore, of three series of equal 



176 



MISS JOAN B. PROCTER ON THE 



and alternating spots (text-fig. 4, c). Group A can only be 
distinguished from B by the fact that these spots are more square 
than those of the latter. In this type the ground-colour is not 
usually darker at A, as is the case in the preceding types. 

Text-figure 5. 





^fMiiaiCTjMlSIWiS'.-ii 





Further development of this type is shown on. a specimen from 
St. Lucia (Cat. p. 536, spec. v). On this snake both the dorso- 
lateral and lateral series are absent, or but faintly marked, the 
markings thus consisting of two squarish spots — A 1, 1 — followed 
by one roundish spot— B 1 — all of which ai^e for' the most part 
confluent with those of the opposite side. 



VARIATION OF THE PIT-VIPER. 177 

The fourth type — L. jararacussu (text-fig. 5) — constitutes the 
third branch in the evolution of the markings. A 1, 5, 2, 6, and 3 
become confluent, forming a subcrescentic marking, in the middle 
of which A 4 may be present or not. All the primitive spots of 
group B unite together to form a dark triangular area, which in 
the course of evolution becomes confluent with group A — i. e., 
the apex of A with the base of B — along the dorsal line. At this 
point the markings darken, and the usual brownish, greyish, or 
olive ground-colour becomes a bright clear yellow. Finally, more 
and more black pigment becomes deposited at A and B, until the 
markings have usurped the place of the ground-colovir, even the 
crescentic marking (A) being partially filled in with black. The 
general effect of Type IV. is therefore a black snake, with yellow 
forked markings issuing transversely from the yellow ventrals, 
which are checkered with black. The dorsal surface of the head 
is completely black, owing to the development and confluence 
between the streaks of the dorsal and dorso-lateral series. The 
post-ocular streak is separated from this area by a narrow stripe 
of the clear yellow ground-colour, which continues slightly below 
the commissure of the jaws. 

The most striking feature about this type of marking is the 
clear yellow of the interspaces in the doi:sal pattern ; in other 
types this is only found on the ventral surface of the snake, the 
dorsal ground colour always exhibiting a superficial powdering of 
dark pigment. It therefore seems probable that in the course 
of evolution this pigment has all aggregated in the spots of 
Groups A and B, and, as more and more becomes deposited, these 
areas further encroached upon the yellow ground-colour. 

V. Phylogenetic Relations between the Types. 

In tracing the evolution of markings from one type to another, 
I have assumed that the anterior parts represent the primitive 
pattern, as in lizards and many mammals (ex. Lacerta 'niuralis, 
Equus quagga). It might appear, however, that in X. cdrox this 
is not the case, the more regularly spotted type, ciiaracteristic of 
the West Indies representing the original pattern, from which the 
clustered, annulated, and forked markings of C. and S. American 
forms were derived. On careful consideration I cannot adopt 
this view for the following reasons: — 1. The manner in which 
the pattern of the Jararacussu develops from Type I. into the 
complicated design of Type TV., and develops on each individual 
posteriorly until the tail is a uniform black, cannot be looked upon 
as primitive, and if this elaborate pattern, become simplified, 
is a further development of the clustered type of marking, the 
other forms must also be more advanced. 2, I hold the greater 
number of spots to be the more primitive (and for this assumption 
there is much correlative evidence among other snakes and 
lizards). The clustered pattern is therefore less advanced than 
the more evenly spotted, for in the former groups A and B 

Phoc. Zool. Soc— 1918, No. XII. , 12 






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180 MISS JOAN B. PROCTER ON THE 

consist of 6 spots each, whilst in the latter group A consists of 5 
and B of 4. Besides these two points, I attach importance to 
the fact that the eight primary series of spots never extend 
beyond the head. 

As we have already seen. Types II., III., and IV. are all 
directly evolved from Type I. ; but their relative positions have 
yet to be discussed. Of the stages of development evinced 
between each of these three and the primitive type, many show a 
tendency for the markings to break up into the spotted type 
(Type III.) just before the tail. Nearly all specimens with mark- 
ings ranging from Types I. to II. show this peculiarit}', also all 
forms intermediate between Types I. and III. In the series 
from Types I. to IV. this is the case in intermediate forms, but 
Type IV. itself maintains its direct line of development, the 
black markings encroaching more and more upon the yellow 
ground-colour towards the posterior part of the snake, the end of 
the tail being black. 

It will therefore be seen that phylogenetically the markings 
of Type III. are not to be regarded as so primitive as those of 
Type II., and that Type IV. is the most modified. 

In the next chapter, on classification, varieties will be based 
on these types of markings in conjunction with their geographical 
distribution. 

Having traced the markings of L. atrox fi-om the most 
primitive onwards, it is of some interest to try to ti'ace these 
markings back to the hypothetical four-paired streaks of primitive 
snakes. 

The first stage towards this would be the equal development of 
group B with group A, and the commencement of the fourth or 
lower labial series on the body. The series would then become 
more equalised, until a pattern were reached of 4 rows of equal 
and alternating spots — independent from those on the oppo- 
site side— which would be but the result of the breaking up 
of the primitive streaks. But L. atrox is very far ahead of this 
prototype, and such cases as might appear at first to be an 
approximation to it, prove on " careful consideration to be 
secondary, an instance of deceptive apparent reversion. 

VI. Classification. 

It will be seen from the foregoing that no very definite 
boundaries can be traced between the varieties of L. atrox, 
whether based on structure or on coloration, so complete are the 
links connecting them. I can, however, recognize four principal 
foi'ras based chiefly on markings, more or less in conjunction 
with geographical distribution, which I think afford on the whole 
a more satisfactory basis for classification than the characters of 
scale-structure, etc. 

The first of these forms is L. atrox, var. affinis, which ex- 
hibits the more primitive type of marking (Type I.), viz., the 



VARIATION OF THE PIT- VIPER. 181 

characteristic chevron, or triangle, which, however, varies slightly 
according to localities. 

The most primitive type of marking is not found in the 
W. Indies proper, or south of Colombia and the Guianas, but 
prevails north as far as Mexico, and appears to be constant in 
Trinidad. Specimens of this form have scales with the high short 
keel. In the W. Indies proper a less primitive variation occurs, 
in which the elemental spots are more confluent and the pattern 
less distinct ; the scales may either be of the short high-keeled 
or long low-keeled type. As this variety extends southwards 
the markings, though still undoubtedly of Type I., frequently 
approach those of the Jararaca, or Jararacussu in type, and the 
scaling maj'- conform to either type. Thus the L. atrox and 
L. lanceolatus of Dr. Vital Brazil, which are marked simply with 
dark triangles on the sides, undoubtedly approach the Jararaca 
of Wied in pattern. A pattern of markings intermediate between 
the primitive type of this variety and that of the Jararacussu 
.occurs in W. Ecuador. L. atrooc, var. ajfinis, is the most widely 
ranging of the four varieties (Mexico to Southern Brazil and 
Peru). All forms of it are marked with triangles or chevrons, 
with their apex turned towards the mid-dorsal line. Group A is 
always darker than B, and encloses an ai-ea darker than the sur- 
rounding ground-colour. The ventrals may be uniform cream- 
colour or yellowish, or blotched or speckled with black. In the 
young the elemental spots, when present, are more distinct, a 
certain amount of confluence or fading taking place with age. 

Form 2, L. atrox, \a,r. jararaca, has been considered a distinct 
species since the time of Wied, but as it does not differ from the 
typical L. atro.v in any appreciable way, save in the dorsal mark- 
ings, I consider it to be a variety only. The Jararaca inhabits 
Brazil, and is especially prevalent in Bahia, Sao Paulo, and 
Rio. It is recognized by its annulated form of markings, pre- 
viously described a^ Type II., consisting of dark, darker-edged 
bands, narrower on the mid-dorsal line than on the sides. These 
bands are more than their own width apart, the interspaces being 
lighter and unspotted. The ventrals may be uniform, spotted, or 
powdered with black. 

Figures of the Jararaca are given in Wied, Abbild. Nat. Bras., 
under the name of Cophias atrox ; Wagler in Spix, Serp. Bras, 
pis. xix., xxi., under the names of Bothrops megcera, B. leuco- 
stigma, B. tessellatus, and B. tceniatus ; and in Jan, Icon. Oph. 
xlvii. pi. iii. under B. atrox. 

A typical specimen of this variety could not be confounded with 
the pi'eceding, but in the British Museum collection there are 
some young specimens from Berbice (Cat. Sn. p. 539, spec, e-k), 
which are exactly intermediate between these three varieties. 

Form 3, tlie typical L. atrox > L. lanceolatus) occurs principally 
in the West Indies, which is the acknowledged home of the 
Fer-de-lance, but also in the Guianas, Venezuela, and curiously 
in Peru, in a somewhat modified form. The markings of this 



182 ON THE VARIATION OF THE PIT-VIPER. 

variety are of the spotted type, described as Type III., consisting 
of three longitudinal series of subequal and alternating dark spots, 
moi-e or less uniformly distributed on a lighter ground-colour ; 
the scales are usually of the low, long-keeled type. In the 
modified form from Venezuela, Berbice, and Peru the spots of 
group A are darker than those of group B, the whole pattern 
thus having a less even appearance. In some cases the mai-kings 
are very indistinct, only the principal spots of group A (A 1, 1, 
2, 3) being discernible. The scales may be of either type. 

Form 4, L. atrox, vsiV. janiracussa, is the distinct colour variety, 
which Lacerda regarded as a valid species in 1884, and which is 
still maintaind as such by Dr. V. Brazil. There is, however, no 
difference between the Jararacussu and L. atro.v except that of 
colour and markings. 

The Jararacussu inhabits Brazil, Sao Paulo, and Rio, and is 
distinguished from the preceding forms by its peculiar markings 
described above as Type IV., and the fact that the interspaces in 
the black areas are bright yellow instead of the brown, grey, or 
olive shades of all the other varieties. The pattern appears to 
consist of transverse yellow forks issuing from the yellov/ ventrals, 
which are blotched with black. The tail is black. The head of 
the Jararacussu is extremely chai'acteristic, the dorsal surface 
being of a uniform black, from which the black post-ocular 
streaks are separated by a narrow streak of yellow, continued 
obliquely from the yellow tliroat. A coloured plate of this variety 
is given in Dr. Brazil's ' Defense contre I'Ophidisme' (2nd edition), 
pi. 14, under the name of Lachesis jararacussu. It is also 
represented in Jan's Iconographie, 47me livr., pi. ii. fig. 3, as 
Bothrops atrox. 



ON DEATHS IN THE SOCIETY S GARDENS. 



183 



9. Report on Deaths of Animals in the Gardens in 1917. 
By J. A. Murray, M.D., B.Sc, F.Z.S., Director, 
Imperial Cancer Research Fund, Pathologist to the 
Society. 

[Received April 23, 1918 : Read April 23, 1918.] 

The accompanying tabular statement of the numbers of deaths 
of mammals, birds, and reptiles (including amphibia and fishes) 
in 1917, shows only minor difierences from the figures recorded 
in 1916 by Professor Plimmer or from the average numbers in 
the previous 5 years. In 1917 rather less than 25 per cent, of 
tlie animals dying had been less than 6 months in the Gardens, 



Table I. 



! 

Mammals, Bieds. 


Reptiles. 




613 1940 
215 745 


446 
368 


In Gardens, l.i. 17. 
Admitted in 1917. 

Total. 

Under 6 months, y,- , 
Over 6 months. "^^'^• 

Per cent, of total. 

iu 1916. 
in 1911-15. 


828 2685 
167=235 i 461=°^* 


814 
80 ^* 


28-4 20-6 


17-4 


28-3 23-4 
27-0 23-3 


28-6 
31-2 



As has been pointed out by Dr. Chalmers Mitchell, little statistical 
value attaches to such tabular statements of combined mortality 
of species with widely difi:erent liability to disease and duration 
of life. The only conclusion to be drawn is that there lias been 
freedom from severe epidemics during the year. 

Table 11. summarises the results recorded under the more 
important causes of death for the chief mammalian orders, and 
for mammals, birds, and reptiles generally. These have been 
compiled from the careful records made by Professor Plimmer as 
regards the first 7 months of the year. There was an interval 
of nearly two months before I entered upon the duties of 
pathologist, and the incomplete data recorded in this interregnum 



184 



DR. J. A. MURRAY ON 

Table II. 





Mammals. 


BiBDS. 


Reptiles. 


"in 


d 

o 

> 
o 


_5 

O 


i 


Edentata. 
• Cheiroptera. 


.2 
■5. 


Total. 


1. General Diseases. 

Tuberculosis 


6 

1 
1 
1 

1 


2 

1 

1 

1 

13 

2 

1 
2 


2 

"2 

26 


3 
1 




1 

i 

"7 

i 

1 

1 

1 

"4 


13 
4 
1 
5 
6 

1 

1 

84 
2 

1 
3 

3 

7 

22 

1 

3 

2 

1 

17 

2 

1 

3 

2 

3 

11 

37 


46 

3 

1 

"2 

197 
5 

3 

1 

88 
7 
2 

58 

1 '2 

i 1 
i 7 

i 1 
I 10 
i 19 

1 81 


4 
1 
9 

... 

"2 
3 

. 
22 




1 --. 


1 Abscess 


Peritonitis 


2 

1 

9 

'3 

i 

i 

2 

3 


1 1 . 


Cestodes 




1 

27 

1 
1 
4 


Ascarides 

HaBmogregarines 

2. Diseases of Bespiraiory 
Organs. 
Atelectasis 


"l 

... 

1 
... 


... 
2 

... 


Pleurisy 

3. Disease.i of Heart. 

Degeneration of Heart-muscle. 
Valvular disease 

4. Diseases of Liver. 
Hepatitis 

5. Diseases of Alimentary Trad. 




3 , 1 

8 1 9 

i ". 

8 2 

1 ... 

1 

2 ... 

1 ' ... 

2 ' ... 

3 3 

9 13 






4 ■ 




1 


Intestinal obstruction 

Intussusception 

Prolapse of Rectum 

6. Diseases of Urinary and 

Generative Organs. 

Nephritis 

Salpingitis 

7. Various. 


1 
... 

4 

1 


i ■■.- 

j 

4 




; 


1 ... 

'2 : ::: 




Haemorrhage 

Injuries discovered post-l 

mortem ) 

Starvation and malnutrition .. 
Killed by companions, rats, &c 


1 

1 

3 

11 


! 12 

1 
i 1 




! 78 






i 


1 


1 





are partly responsible for swelling the totals under the heading 
"not diagnosed " in Table II. 

Protozoan Parasites. — In addition to the three reptilian deaths 
in Table II. ascribed to Hsemogregariiies, these parasites have 
been observed in a number of snakes dying from other causes. 



DEATHS IN THE SOCIETY'S GARDENS. 185 

They have revealed an apparently definite dimorphism of the 
schizonts in the circulating blood, a long slender form which 
produces great deformation of the red cell, the hsemoglobin being 
retained, and a short stout form which produces much less 
deformity but exhausts the haemoglobin very rapidly. The 
significance of the two forms and their relation to the schizogony 
in the liver-cells remain to be determined by further study, for 
which a considerable material has been preserved. 

Malignant neiv Groivths. — In two mammals death was ascribed 
to carcinoma and in two birds to sarcoma. Of these one case of 
squamous-cell carcinoma of the fauces and palate of a Dingo, 
probably arising from the epithelial covering of the tonsil, has 
been examined personally. In addition a new growth was found 
on examination of a Leopard which died with bronchiectatic 
cavities in the lungs. The growth in this case pi'oved to be an 
adeno- carcinoma lying on the ventral surface of the trachea in 
the thorax. The tissue of origin covild not be determined. 

(Two further cases have been observed in 1918: a carcinoma 
of the liver in a Marsh-Buck and a teratoma of the testis in a 
Golden Eagle.) 

Comparative Pathology of the Thyroid. — The special value of 
the material occurring in the Prosectorium consists in the 
opportunity offered for a comparative study of pathological and 
physiological problems which are of general interest, a.nd special 
attention has been directed to the thyroid gland. The thyroid 
occurs throughout the -whole vertebrate series and, with the 
exception of the cyclostome fishes, presents the same histological 
picture in all. From studies in man and mammals in various 
pathological conditions an important role has been asci'ibed to it 
in the reaction of the body to a variety of intoxications. The 
results of physiological experiment indicate other important 
functions both in health and disease. It was therefore of interest 
to note the contrast in the appearance of the gland in cold- and 
warm-blooded animals respectively in severe infections. In 
warm-blooded animals dying under these conditions extreme 
congestion of the whole gland is piactically constant. ISTothing 
of the kind has been encountered in the reptiles examined, 
although a large projDortion presented severe septiceemic con- 
ditions after death. This result is unfavourable to the view that 
the thyroid plays- the pai't of a neutraliser of toxic substances in 
the body. It is in much better harmony with the view that the 
changes in the thyroid in these conditions are the expression of 
its participation in the heat-regulating mechanism of the body. 
In poikilothermic animals one would expect these changes to be 
absent. 

E^ffect of Diet on the Thyroid. — Although attention has been 
drawn to the effect of a meat diet on the thyroid, no observations 
are recorded on the results of the natural experiment presented 
by the occurrence of meat-eating and plant-eating groups in birds 
and mammals. The analysis of the observations in birds presents 



186 ON DEATHS IN THE SOCIETY'S GARDENS. 

many difficulties, and will be undertaken later. In mammals, 
liowever, tlie comparison is much easier, and the preliminary 
survey which has been carried out shows that in Oarnivora the 
thyroid is from two to three times as large as in Ungidata of 
corresponding body-weight. The key to the problem is probably 
furnished by Marine and Lenhart's work on thyroid enlargement 
in tiout kept in hatcheries. These authors have shown that the 
goitrous condition, which so frequently supervenes, is due to the 
diet of horseflesh with which the fish are fed and that it can be 
obviated and even cured l)y supplying a ration of sea-fish. Their 
investigations have shown that the constituent deficient in the 
horseflesh diet is iodine. The necessity for fixing the inadequate 
amounts- present in flesh is met by a hypertrophy of the gland. 
"When adequate amounts of iodine are supplied either in the form 
of sea-fish or soluble iodides, the enlar£;ement subsides. 



THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 187 

EXHIBITIONS AND NOTICES. 

February 5th, 1918. 

Dr, A. Smith Woodward, F.R.S., Vice-President, 
in the Oliair. 

The Secretary read the following Report on the Additions 
made to the Societ3''s Menagerie during the months of November 
and December, 1917 : — 

November. 

The registered additions to the Society's Menagerie during tlie 
month of November were 38 in number. Of these 28 were 
.acquired by presentation an<l 10 were received on deposit. 

The number of departures during the same period, by death 
and removals, was 88. 

Amongst the additions special attention may be directed 
to:— 

1 Brindled Gnu [Gorgon taurinus) $ , from South Africa, 
deposited on November 21st. 

1 Wilson's Bird of Paradise [Schlegelia loilsoni), from Waigiou, 
and 2 Himalayan Goldfinches [Uard-uelis caniceps), pi'esented by 
Alfred Ezra, F.Z.S. 

1 Lesser Vasa Parrot [Coracopsis nigra), from Madagascar, 
presented by The Marquess of Tavistock, F.Z.S. 

December. 

The registered additions to the Society's Menagerie during the 
month of December were 17 in number. Of these 7 were 
acquired by presentation and 10 were received on deposit. 

The number of departures during the same period, by death 
and removals, Avas 81. 

Amongst the additions special ;ittention may be directed 
to:— 

1 Grey Seal {Halichoervj% grypns), from the North Sea, presented 
by T. Witherwick on December 24th. 

1 European Flamingo {Ph(j(:nico2)terus roseus), horn S. Europe; 
2 South- Amei'ican Flamingoes {Ph. chilensis), from S. America ; 
2 Black-necked Swans {Cygnas melaiiocephalus) find 2 Coscovoba 
Swans [C'oscoroba coscoroba), from S. America : deposited on 
December 31st. 



Mr. D. Seth Smith, F.Z.S., Curator of Birds, exhibited and 
remarked on a series of lantern-slides made from photogi^aphs 



188 MR. T. E. WHITEHEAD ON THE WILD DINGO. 

of Reptiles taken in the Gardens, directing special attention 
to those showing feeding habits of the Black Ciibo {Oxyrhopus 
cl(jelia), which attacks and devours poisonous snakes. 



The Wild Dingo (Canis familiaris, var.). 

The Secretary read the following letter from 'Mr. Thomas E. 
Whitehead : — 

" During a recent visit to England, I spent a delightful day at 
the Zoological Gardens, but was rather surprised at the animals 
exhibited as Dingos. Now, Sir, I spent a great many years as a 
stockman and drover in the back country of South Australia, 
New South Wales, Queensland, and Northern Territory, besides 
travelling and prospecting over a large part of Western Australia 
during the past 23 years, but I have never met with a Dingo 
anything like those I saw at the London Zoo. 

" I have enquired from my friends if they could tell me what 
part of Australia they could have come from, and I am told that 
many years ago a Mastiff got away with the Dingos about Mount 
Kosciusko and crossed the breed in that district. So probably 
that is where your Dingos came from. They are too tall, too 
long, too thick muzzled, and too short-haired to be true Dingos. 
The true Dingo resembles the fox a great deal, but is larger, his 
nose is pointed, forehead fairly broad, has longish hair, and a tail 
almost as bushy as the fox. There are two colours, according to 
the colour of the country he inhabits. Those in the Tableland 
and Mountainous districts are usually of a sand}^ brown colour, 
while those in the yellow sand-hill country are usually sandy 
yellow, and have longer hair than those on the Tablelands. I 
once saw a pack of about a dozen of the darker kind in the fore- 
noon on the south side of Lake Yandandaninna, and towards 
evening I saw a pack of the lighter colour on the noith side of 
the lake ; they had come for a dr-ink from the sandhills to the 
northward. 

i' Dingos never bark, neither do they learn to bark when 
tamed. The Bushman can always distinguish between the howl 
of a Dingo and that of a Domestic Dog. Like the fox, he is 
very partial to poultry, and is a terror to poultry-owners in the 
new townships. He will kill sheep by the dozen just for the 
fun of doing so. If hungry, he will rip a hole in the flank and 
eat his fill of the liver and leave the rest. I once heard sheep 
rushing on a bright moonlight night. On going to investigate I 
saw our Overseer's prize Kelpie rounding up about 300 sheep, 
while in the centre was a Dingo slut enjoying herself by killing 
sheep wholesale. I fired my rifie at the Kelpie but missed, then 
fired at the slut, and next morning tracked her up by blood 
marks and found her nearly dead, so quickly despatched her. 



ON A NEW SPECIES OF THE GOLDEN MOLE. 189 

She had killed 11 sheep and had bitten many more, some of 
them dying during the next few days. 

" Thos. E. Whitehead, 

262 Bulwer Street, 

North Perth, 
Western Australia." 



On a New and a Rare Species of the Golden Mole 
(Bematiscus). 

Dr. R. Broom, O.M.Z.S., exhibited specimens of two rare 
South-African Golden Moles, and made the following re- 
marks : — 

Hitherto the giant moles from the eastern part of Cape 
Colony have been referred to Bematiscus t7-evelyani, but it is 
quite manifest that there are at least two species. The specimen 
which I make the type of this new species was sent to me by 
, Miss Ivy Lesch fi'om St. Cuthberts, Isolo, and as it diflfers very 
markedly from Bematiscus trevelyani of the Pirie Bush or any of 
the hitherto described species, I have much pleasure in naming it 
after the discoverer, to whose interest in natural history we also 
owe Bematiscus transvaalensis. 

Bematiscus leschce is a moderately large mole, probably about 
180 mm. in length, but as I have only the dried skin, it is 
impossible to be quite sure of the length. It is manifestly con- 
siderably smaller than B. trevelyani. The fur is of a fairly 
uniform dark brown — lighter on the abdomen. On the lips the 
short fur is somewhat reddish brown. On the upper side of the 
head the fur is^also rather lighter than furt))er back, owing to 
the short under-fur being pale reddish brown. On the back and 
greater part of the body the fur is about 18 mm. in length— a 
dark slaty very tine under-fur With coarse-tipped hairs whose tips 
of 7 mm. overlap the under-fur. The tips have the last 2 or 
3 mm. dark brown and the more proximal | light brown. 

The following are the measurements, of the claws of the front 
foot: — 1st 2-5 mm., 2nd 6 mm., 3rd 11 mm., 4th 1-5 mm. 

The following are the principal measurements of the skull : — 

mm. 

Greatest length 35*6 

Greatest width 21-4 

Greatest height 17 

Width between orbits 7'5 

Dental series 14 

Molar series 8 

Width across the premolars 1 '8 

Width between last premolars 4-6 

The skull has the large bulla for the malleus well projecting 
as in B. villosa and B. traihsvaalensis, and the affinities are much 
closer with these two species than with B. trevelyani. The mole 



1 90 THE SECRETARY OX ADDITIONS TO THE MENAWERIE. 

f lom Maritzburg described by Dobson, and doubtfully referred by 
bim to B. villosa is, I think, undoubtedly a distinct species, and 
may be called Beinatiscus dobsoni. 

To the genus Bematiscus there would thus be refeired five 
known species — B. villosa and B. dobsoni. from Natal, B. trans- 
vacdensis from the Transvaal, and B. trevelyani and B. leschce 
from Eastern Cape Colony. 

The second exhibit is a specimen of the rare Golden Mole — 
Chrysochloris sclateri. Originally discovered in the Nieuwveld, 
north of Beaufort West, and napied in honour of Mr. W. L. 
Sclater, it was next discovered at Morija in Basutoland, ?>bO miles 
from the oi'iginal locality. In an unexplored country a species 
occurring at two such remote localites would not excite any 
wonder, but in South Africa our researches on the fauna have 
gone so far that we can say not only that the species has never 
been found in the intermediate area, but that no Golden Moles 
inhabit the northern part of Cape Colony. Six years ago I 
discovered tracks of Chrysochloris on Compass Berg, and I have 
since been fortunate, through the kindness of Miss K. Jansen, of 
Wilgebosch, New Bethesda, in getting a specimen which proves 
to be also C. sclateri. This discovery brings the known localities 
130 miles nearer, and suggests that the species may originally 
have extended to Basutoland by the Winterberg and Stormberg. 
Possibly it may yet occur in these localities. The black mole of 
the south coast, which I named C. duthiece, is an extremely closely 
allied species. It occurs at Knysna, and possibly extends east- 
wards to Port Elizabeth. C. sclateri with its 40 teeth is probably 
near the ancestral form from which C. hottentota and allied forms 
with 36 teeth are derived. 



February 19th, 1918. 

Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following Report on the Additions 
made to the Society's Menagerie during the month of January 
1918:— 

The registered additions to the Society's Menagerie during the 
month of January were 26 in number. Of these 16 were 
acquired by presentation, 8 wej'e received on deposit, and 2 by 
purchase. 

The number of departures during the same period, by death 
and removals, was 80. 

Amongst the additions special attention may be directed to : — 

1 White-nosed Coati {N'asua narica), from Tampico, presented 
by H. C. Banbury on January 19th. 



AN AFRICAX CTVET ATTACKING HUMAN BEINGS. 191 

1 Otter (^fjutra Intra), from Hampton Court, presenterl by 
H. Tagg on January 18th. 

2 8traw-neckecl Ibises (Carphibis spinicollis), from Australia ; 
1 Scarlet Iliis {Eudocimus ruber), from Para ; 1 Himalayan 
Monaul {Lophophoras iinpejjanics) J , from Himalayas ; 1 Peacock 
Pheasant [Poh/plectrum chinqids) S , from Burma.h ; 2 Swiuhoe's 
Pheasants {Gennceus sioinhoii) (S $ , from Formosa, presented by 
Major The Hon. Waldorf Astor, F.Z.S. 



Dr. SairiH Woodward, F.R.S., V.P.Z.S., exhibited a copy of 
an incised drawing of a hunted deer, pierced by arrows, made by 
Palaeolithic man in the cave of La Peiia, San Roman de Oandamo, 
Asturias, Spain. It was lately published by Dr. Hernandez- 
Pacheco in no. 17 of .the memoii's of the Spanish commission on 
prehistoric investigations. 



An African Civet attacking tluman Beings. 

The following letter, communicated by Professor Poulton, 
F.R.S., F.Z.S., was read from Captain G. D. Hale Carpenter, 
M.D., giving an account of a case which had come under his 
personal observation in which an African Civet attacked human 
beings. 

"On the night of Dec. 10-1 1th, 1917, an Indian and an 
African, employees of the railway, were sleeping in the verandah 
of the station building when the latter was awakened by a bite 
on his toe, and' found to his alarm what he thought was a young 
leopard — -it was a very dark night and without a moon. It 
viciously attacked the two men, but they managed to catch it 
and throw it down a well, as they had no stick or other weapons 
handy.: they both came to hospital in the morning to have their 
wounds dressed — the African had a contused and punctui-ed 
wound on the ball of the great toe : the forearm of the Indian 
was superficially lacerated. When they brought up the body of 
the culprit, which I had decided in my mind would prove to be a 
Serval cat, as I thought it unlikely the two men would without 
weapons have overpowered a Leopard cub. But to my astonish- 
ment the draggled carcase was that of a rather small, old. Civet ! 

" The men said there had been two of them. 

" I should think undoubted instances of unprovoked assault by 
a Civet on mankind must be rare. 

" G. D. Hale Carpenter, M.D., 

Capt, Uganda Med. Service." 

" On the Central Railway of ex-German East Africa, 
17 miles W. of Tabora, 
December 12, 1917." 



192 A CH^TODONT FISH WITH ARABIC CHARACTERS. 

Professor E. W. MacBride, M.A., D.Sc, F.R.8., F.Z.S., gave 
an account, illusti-ateil by lantern-slides, of his recent investi- 
gations into the development of the Sea- Urchin (^Echinocardmm 
cordatum). 



March 5th, 1918. 

Dr. A. Smith Woodward, F.E„S., Vice-President, 
in the Chair, 

Mr. D. Seth Smith, F.Z.S., exhibited skins of the Hoatzin 

{Opisthocomus cristatus), and described the habits and distribu- 
tion of the species, illustrating his remarks with lantern-slides. 



A Chcetodont Fish ivith Markings resembling Arabic 
Characters. 

Mr. C. Tate PtEOAN, M.A., F.R.S., F.Z.S., exhibited photo- 
graphs of an Indo-Pacific Chsetodont Fish [Holacanthus semi- 
circulatus Cuv. et Yal.). Two of these had been sent to him 
by Major H. R. Cartwright, Commandant of Police, Zanzibar ; 

Text-figure 1 . 



they were of a specimen that had been sold in the fish-market 
for a penny ; the man who bought it was going to eat it and cut 
off the tail and threw it on the ground ; another man picked it 
up and called out that it had writing on it, and, indeed, on one 
side of the caudal fin was written in old Arabic characters 
" Laillaha Illalah " (There is no God but Aljfih) and on the 
other side " Shani-AUah " (A warning sent from Allah). The 



THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 193 

news caused great excitement in the market place ; the fish 
changed hands at rapidly increasing piices until 5000 rupees was 
•ofiered ; tlie fish was regarded as sacred and Major Cartwright, 
who put it in formalin for the owner and had it photographed, 
thought it might become the object of pilgrimages. 

The other photographs exhibited by Mr. Regan were of ex- 
amples of this species in the collection of the British Museum 
(Natural History), and illustrated the changes in coloration that 
take place during growth. In young specimens the ground- 
colour is almost black, the body is crossed by a few curved white 
stripes, and the posterior half of the caudal fin is clear. In larger 
fish 3 or 4 inches long the general ground-colour is paler, but is 
•dark-spotted ; the stripes have increased in number by the addi- 
tion of narrow ones between the original ones, and still narrower 
ones between these, so that there may be as many as 24 stripes 
instead of the original 6; also the posterior , part of the tail is 
now darked and is crossed by 3 pale stripes, of which the posterior 
2 may be discontinuous or connected or replaced by longitudinal 
bars ; it is these which may simulate Arabic character's. This 
stage was described as Ilolacanthits alternans Cuv. & Val,, and 
from now onwards the ground-colour becomes paler and the dark 
spots better defined, the pale stripes disappear anteriorly, and 
posteriorly still increase in number but break up into spots and 
vermiculations, so that a fish of seven inches long has quite a 
difi'erent appearance, and indeed was desciibed by Bleeker as 
a distinct species, H. lepidolepis. 



March 19th, 1918. 

Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following report on the Additions 
made to the Society's Menagerie during the month of February 
1918:— 

The registered additions to the Society's Menagei'ie dui-iug 
the month of February were 19 in number. Of these 14 were 
acquired by presentation, and 5 were received on deposit. 

The number of departures during the same period, by death 
and removals, was 71. 

Amongst the additions special attention may be directed 
to:— 

2 Stanley Cranes {Tetrapteryx pm'adisea), from South Afi'ica, 
and one Black-necked Crowned Crane [Balearica 2^ct'>^onina), 
from West Africa, presented by W. H. St. Quintin, F.Z.S., on 
Februaiy 21st. 

1 American Bison {Bison amei'icanus), from N. America, 
deposited on February 15th. 

Proc. Zool. Soc— 1918, No. XIII. 13 



194 ON FOSSIL HOSTRAL TEETH OF SAW-FISHES. 



April 9th, 1918. 

Dr. A. Smith Woodward, F.E.S., Vice-President, 
in the Cliaii-. 

The Secretary read the following Report on the Additions, 
made to the Society's Menasrerie during the month of March 
1918: — 

The registered additions to the Society's Menagerie during the 
month of March were 31 in number. Of these 23 were acquired 
by presentation, 6 were received on deposit, 1 by purchase, and 
1 in exchange. 

The number of departures dui-ing the same period, by death 
and removals, was 93. 

Amongst the additions special attention may be directed 
to :— 

1 Jungle-Cat {Fells chaiis) .$ , fi'om India, purchased on 
March 9tJi. 

2 Hog-Deer {Axis porcimcs), S 2 > from India, deposited on 
March 5th. 

IKing Penguin {Aptenodytes patagonica), from South Georgiay 
received in exchang-e on March 19th, 



Miss L. E. Cheesman, Assistant Curator of Insects, exhibited 
specimens of an East- African homopterous insect, Ityrcea nigro- 
cincta, sent by the Curator of the Museum at Nairobe. 

Special interest is attached to these insects by reason of their 
habit of clusteiing in colonies on a plant-stem in imitation of the 
inflorescence. There is a coloured form and a green form of 
the adult insect, and when the latter are found clustering on the 
apex of the flower-spike, they bear an extraordinary resemblance 
to unopened buds. 

An interesting account of a similar species was published by 
Dr. Gregory in his book ' The Great Rift Yallej,' with a sketch 
of the insects. They have also been figured in a paper read by 
Mr. Hinde before the Entomological Society in 1902. 



Dr. A. .Smith Woodward, V.P.Z.S,, exhibited fossil rostral 
teeth of Eopristis smd Pristis, and referred to the progressive 
changes in the rostral teeth of the Pristidae, or saw-fishes, dinging 
geological time. In the Cretaceous genus, Sclerorhynchus, these 
teeth only differ from the ordinary shagreen in their enlargement 
and the elongation of their apical portion. Their pulp-cavity is 
restricted to the basal half, and the distal portion is traversed by 
several very i irregular lai-ge vascular canals, mainly longitudinal 
in direction. In the genus Eopristis, not known later than the 



ON FUR SALES IN THE UNITED STATES. 195 

Eocene period, the teeth are still loosely ranged along the edge of 
the rostrum, not in deep sockets ; but their structure is nearly 
that of Pristis, the vascular canals being only much more iri'egular 
and not yet clearly the centres of well-marked hexagonal prisms. 
Pristis itself, with the rostral teeth in deep sockets, and with the 
well-known prismatic structure, ranges from the Eocene period 
onwards. 



Mr. G. A. BouLENGER, F.R.S., exhibited the head of a Hydro- 
cyon goliath Blgr., from the Congo, a fish attaining a length of 
four feet, the largest and most formidable representative of a 
genus of the family Oharacinidas inhabiting the pi'incipal rivers 
of Africa. The Nile species is known as the Kelp-el-Bahr, or 
River-dog, of the Ai-a,bs, and a third species, common in the 
Zambesi and Limpopo systems, is called Tiger-fish by the 
colonists. 

The object of this exhibition was to show the enormous shark- 
like teeth to which special interest attaches owing to a recent 
publication by Dr. Eastman, who has pointed out their close 
similarity to the fossils known as Onchosaurus Gervais (originally 
referred to the Mosasaurs), Ischyrhiza Leidy, and Gigantichthys 
Dames, which appeal's to indicate the existence of the Characinidse 
as far back as the Upper Cretaceous, a lange in time which 
Mr. Boidenger had predicted as probable thirteen years ago, and 
which is of special importance in explaining the present distri- 
bution of this family*. 



April 23rd, 1918. 

A. Ezra, Esq., 7ice-President, in the Chair. 

Mr. D. Seth Smith, Curator of Birds, exhibited and made 
remarks on a Zulu Head-dress made of the plumes of the male 
Long-tailed Whydah, Chere progne. 



The Secretary called attention to an advertisement that had 
recently appeared in the London Press, announcing Fur Sales by 
Public Auction about to take 'place in the United States. The 
sales in question were only examples of what took place annually 
in London and other important commercial centres. The numbers 
advertised Avere smaller than usual, no doubt on account of 
the War, but they included very large quantities of animals the 

* Cf. C. R. Ac. Sci. Paris, clxvi. 1918, p. 197. 



196 ON SPECIMENS AFFECTED WITH RICKETS. 

extinction of which could not be far distant, unless measures 
were carried out to protect them. In the opinion of the speaker, 
which was confirmed by the Meeting, there was urgent need for 
drastic measures to protect Mammals. The protection of Birds 
appealed to popular sentiment, and was zealously advocated by 
many influential organisations. The danger that threatened 
Mammals was even greater, and, on account of their higher 
intelligence and more sensitive nervous organisation, the cruelty 
involved in the methods of hunting, trapping, and killing them 
was incomparably greater. 



Professor Wood-Jones, F.Z.S., Honorary Acting Prosector, 
exhibited and made remarks on specimens from the Prosectorium 
illustrating the eflfects of Rickets. He also exhibited a set of 
anatomical preparations useful for teaching purposes, made from 
material obtained from the Society's Collection. 



ABSTRACT OF THE PROCEEDINGS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDOiN.' 

February 5tb, 1918. 

Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmcil. 

The Secretary read a Report on the Additions to the 
Menngerie in the months of jSTovember and December, 1917. 

Mr. D. Seth Sivrixn, F.Z.S., Curator of Birds, exhibited and 
made remarks on a series of lantern-slides made from photo- 
graphs of Reptiles taken in the Gardens, directing special 
attention to those showing feeding habits of the Black Ciibo 
{Oxyrhopus cloelia), which attacks and devours poisonous snakes. 

The Secretary read a letter from Mr. Thos. E. Whitehead 
containing observations on the Avild Dingo of Australia. 

A paper by Professor B. L. Buatia and Baini ^Prasiiad 
(communicated by Lieut.-Colonel J. Stephenson, D.Sc, I. M.S., 
F.Z.S.), entitled the " Skull of liana tigrina Daud., was read. 

Mr. G. A. BouLBNGER, F.R.S., F.Z.S., presented his paper 



entitled " Descrij)tion of a 
from Upper Bui-ma." 



new Snake of the genus Oligodon, 



* This Abstract is piiblislied by the Society at its offices, Zoological Grardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of 8ix Shillings per annum, payable in advance. 



2 

Dr. R. Broom, O.M.Z.S., exhibited specimens of two rare 
South African Golden Moles. The one was described as a new 
species of Beniatismos, B. leschce. Hitherto the Giant Moles of 
the Eastern Cape Colony have been referred to B. trevelT/ani, hut 
the present type from St. Cathberts, Isolo, differs from B. tre- 
velyani and agrees with B. transvacdensis and B. villosa in having 
the temporal bulla markedly projecting from the side of the 
skull. The skull measures 35-6 mm. in length and 21-4 mm. in 
width, being thus veiy appreciably smaller than B. trevelyani. 

The other specimen exhibited was one of the rare mole 
Chrysochloris sdateri. Hitherto it has been only known from 
the Nieuwveld and from Basutoland — localities 350 miles apart. 
The present specimen was from New Bethesda, 130 miles nearer 
to Basutoland than the original locality. 



The next Meeting of the Society for Scientific Business will be 
held on Tuesday, February 19th, 1918, at 5.3Q p.m., when the 
following communications will be made : — 

Professor E. W. MacBride, M.A„ D.Sc, F.R.S., F.Z.S. 



The development of Echinocardmm cordatttm^ 
Captain G. D. Hale Carpenter, ¥I.D. 



An African Civet attacking Human Beings. (Communi- 
cated by Pi*ofessor Poulton, F.B.S,) 

L. A. Lantz (Moscow). 

Reptiles from the River Tajan. (Communicated by G. A. 
Boulenger, F.R.S., F.Z„S.) 



The following papers have been received : — 

Miss Maude L. W. Cleghorn, F.Z.S., F.L.S., F.E.S. 

First Report on the Inheritance of Visible and Invisible 
Characters iu Silkworms, 



3 

R. I. PococK, F.R.S., F.Z.S. 

On the External Characters of the Lemurs and Tarsius. 

Dr. R. W. Shufeldt, C.M.Z.S. 

Notes on the Osteology of the Young of the Hoatzin 
{Opisthocomus cristatus) and other points on its Morphology. 



Comnmnications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society op London, 
• Regent's Park, London, N.W. 8. 
Fehruari/ I2th, 1918. 



No. 176. 



ABSTRACT OF THE PROCEEDINGS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON,' 

February 19th, 1918. 

Dr. A. Smith Woodward, F.R.S., Yice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the Additions to the 
Menagerie in the month of January, 1918. 

Dr. Smith Woodward, F.R.S,, V.P.Z.S., exhibited a copy of 
an incised drawing of a hunted deer, pierced by arrows, made by 
Palaeolithic man in the cave of La Peiia, San Roman de Candamo, 
Asturias, Spain. ^It was lately published by Dr. Herndndez- 
Pacheco in no. 1 7 of the memoirs of the Spanish commission on 
prehistoric investigations. 

A letter, communicated by Professor Poulton, F.R.S,, F.Z.S., 
was read from Captain G. D. Hale Ca,rpenter, M.D., giving an 
account of a case which had come under his personal observation 
in which an African Civet attacked human beings. 



A paper by L. A. Lantz of Moscow, communicated by Mr. G. 
A. BouLENGER, F.R.S. , F.Z.S. , describing a collection of reptiles 
made in Transcaspia and now in the Zoological Museum of 
Moscow University, was read. 



* This Abstract is published by the Society at its ofSces, Zoological Gardens, 
Eegent's Park, N.W., ou the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may bo obtained on the 
day of publication at the price of SixjJence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable iu advance. 



6 

Professor E. W. MacBride, M.A., D.Sc, F.R.S., F.Z.S., gave 
an account, illustrated by lantern-slides, of his recent investi- 
gations into the development of the Sea-Urehin (Echinocardnom 
cordatnni). 



The next Meeting of the Society for Scientific Business will be 
held on Tuesday, March 5th, 1918, at 5.30 p.m., when the 
following communications will be made : — 

R. I. PococK, F.R.S., F.Z.S. 

On the External Characters of the Lemurs and Tarsius. 
(Illustrated by lantern-slides.) 

Sir George F. Hampson, B t., F.Z.S. 

A Classification of the Pyrcdidce, Subfamily Hypsotropince. 



The following Paper has been received : — 

Miss Maude L. W. Cleqhorivt, F.Z .S., F.L.S., F .E.S. 

First Report on the Inheritance of Visible and Invisible 
Characters in Silkworms. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society op London, 
Regent's Park, London, K.W. 8. 
February 26th, 1918. 



ABSTRACT OF THE PROCEEDINGS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON/ 

March 5tli, 1918. 

Dr. A. Smith Woodward, F.R.S., Vioe-Presicieiit, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

Mr. D. Seth Smith, F.Z.S., exhibited skins of the Hoatzin 
(^Opisthocomus ci'isiatus), and described the habits and distribu- 
tion of the species, illustrating his remarks with lantern-slides. 

Mr. R. I. PococK, F.R.S., F.Z.S., gave an account of his 
communication entitled " On the External Characters of the 
Lemurs and Ta7'sius." The observations recorded were based, 
except in the cas-e of Tarsius, upon specim-ens that had lived in 
the Society's Gardens. They related chiefly to the muzzle, the 
ear, vibrissae, hands and feet, and the perineal organs. The 
author, in conclusion, stated his opinion that Tarsius should be 
removed from the Lemuroid Friraates and classified with" the 
Monkeys. He proposed to divide the Primates into two primary 
groups, the Strepsirhini for the Lemurs and the Haplorhini for 
Tarsuis and the rest, the Haplorhini being further divided into 
the Tarsibidea for Tarsius a,nd the Pithecoidea for Monkeys, 
Apes, and Man, 

A communication from Sir George F. Hampson, Bt., F.Z.S., 
contained a classification of the Hypsotropince which the Author 
described as a rather obscure group of the Pyralidai, of very 
uniform appearance and differing chiefly in structure. 



* This Abstract is published by the Society at its offices, Zoological Gardeus, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on 
the day of publication at the price of Sixpence, or, if desired, sent post-free 
for the sum of Six SMllinr/s per annum, payable in advance. 



8 

Uv. C. Tate IIegan, M.A., F.R.S., F.Z.H., exliibited pLoto- 
graplis of an Indo-Pacific Cliaitodont Fisli [Uolacanthtis semi- 
circulatus Cuv. et Yal.). Two of these had been sent to hiiu 
by Major H. K, Oa.rtwright, Commandant of Police, Zanzibai^ ; 
they were of a specimen that had been sold in the fish-mai-ket 
for a penny ; the man who bought it was going to eat it and cut 
off the tail and threw it on the ground ; another man picked it 
np and called out that it had writing on it, and indeed on one 
side of the caudal fin was written in old Arabic characters 
"Laillaha Illalah " — There is no God but Allah — and on the 
other side " Shani- Allah " — A warning sent from Allah. The 
news caused great excitement in tlie market place ; the fish 
changed hands at i-apidly increasing prices until 5000 rupees was 
offered ; the fish was regarded as sacred and Major Cartwright, 
Avho put it in formalin for the owner and had it photographed, 
thought it might become the object of pilgrimages. 

The other photographs exhibited by Mr. Regan were of ex- 
amples of this species in the collection of the British Museum 
(Natural History), and illustrated the changes in coloration that 
take place during growth. In young specimens the ground- 
colour is almost black, the body is crossed by a few curved white 
stripes, and the posterior half of the caudal fin is clear. In larger 
fish 3 or 4 inches long the general ground-colour is paler, but is 
dark-spotted ; the stripes have increased in number by the addi- 
tion of narrow ones between the original ones, and still narrower 
ones between these, so that there may be as many as 24 stripes 
instead of the original 6 ; also the posterior part of the tail is 
now darked and is crossed by 3 pale stripes, of which the posterior 
2 may be discontinuous or connected or replaced by longitudinal 
bars ; it is these which may simulate Arabic characters. This 
stage Avas described as Holacanthus cdternans Cuv. & Val., and 
from now onwards the ground-colour becomes paler and the dark 
spots better defined, the pale sti'ipes disappear anterioi-ly, and 
posteriorly still increase in number but break up into spots and 
vermiculations, so that a fish of seven inches long has quite a 
diflerent appearance, and indeed was desciibed by Bleeker as 
a. distinct species, II. lejyidolepis. 



9 

The next Meeting of the Society for Scientific Business will be 
held on Tuesday, March 19th, 1918, at 5.30 p.m., when the 
following- comnumications will be made : — 

Dr. S. F. I-Iamiee, M.A., Sc.D., F.R.S., F.Z.S. 

Notes on Cetacea stranded on the British Coasts during the 
last Five Years. 

I\[iss Maude L. W. Oleghorn, F.Z.S., F.L.S., F.E.S. 

First Report on the Inheritance of Visible and Invisible 
Oharactei'S in Silkworms. 



The following Paper has been received : — 
Dr. Branislav Petronievics. 



Comparison between the Lower Jaws of the Cynodont Rep- 
tiles GomiJiliognailius and Gynognathus. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

— Secretary. 

Zoological Society of London, 

Regent's Park, London, K.W. 8, 
J/arcA 12fA, 1918. 



No. 178. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON.* 

March 19th, 1918. 



Dr. A. Smith Woodward, F.R.S., Yice-Presideut, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the Additions to the 
Menagerie in the month of February, 1918. 

Dr. S. F. Harmer, F.R.S., V.P., gave an account, illustrated 
by lantern-slides, on observations made on Cetacea stranded on 
the British Coast during the last five years. 

A paper by Miss Maude L. W. Cleghorn, F.Z.S., entitled 
" First Report on the Inheritance of Visible and Invisible 
Characters in Silkworms," was communicated. 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Eegent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the 'Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on tha 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



12 

The next Meeting of the Society for Scientific Business will be 
lield on Tuesday, April 9th, 1918, at 5.30 p.m., when the 
following communications will be made : — 

G. A. BouLENGER, LL.D., F.B.S., F.Z.S. 

Exhibition of a head of the Charasinid Fish, Flydrocyon 
goliath. 

Miss J. B. Procter, F.Z.S. 

On the Variation of the Pit-Viper, Lachesis atrox. 



The following Paper has been received : — 
Dr. Bbanislav Petronievics. 

Comparison between the Lower Jaws of the Cynodont 
Reptiles Gompliognathus and Oynognathus. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
Zoological Society of London, 
Regent's Park, London, N.W. 8. 
liarch 2Qth, 1918. 



No. 179. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON, 

April 9tli, 1918. 



Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a Report on the Additions to the 
Menagerie in the month of March, 1918. 

Miss Cheesman, Acting Curator of Insects, exhibited and 
made remarks on examples of the East African Plant-bug, 
Itymcea nigrocincta, sent from the Nairobi Museum. 

Dr. A. Smith Woodward, F.R.S., Y.P.Z.S., exhibited fossil 
rostral teeth of the Sawfishes Eophristis and Fristis, and ref ei-red 
to the progressive changes in the rostral teeth of the Pristidss 
during geological time. 

Mr, G. A. BouLENGER, F.R.S., exhibited the head of an 
example of HydroGyon goliath, Blgi'-j from the Congo, a fish 
attaining the length of four feet. The object of the exhibition 
was to show the enormous shark-like teeth, to which special 
interest attaches, owing to a similarity, recently pointed out by 
Dr. Eastman, to fossil teeth occurring in the Upper Cretaceous, 
which would appear to indicate the existence of Characinidfe in 
that geological epoch, a range in time which Mr. Boulenger had 
predicted as probable thirteen years ago. 

* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Pai-k, N.W., ou the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on tlu^ 
day of publication at the price of Sixjjence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



14 

Mr. G. A. BouLENGER also, in the absence of the author, gave 
an account of a communication by Miss J. Proctor, F.Z.S., 
entitled " On the Variation of the Pit-Viper, Lachesis atrox." 
The paper dealt with the variation of the principal characters of 
the Central and South American Pit-Viper, Lachesis atrooc L., 
of which she regarded L. lanceolatus Lacep. as a synonym, and 
L. affinis Gray, jararaca Wied, and jararacussu Lacerda, as 
varieties. She laid special stress on the different patterns of 
markings, discussing their evolution and regarding that shown 
by the more northern f'orm, L. affinis, as the most primitive, 
from which all others could be derived. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, April 23rd, 1918, at 6.30 p.m., when the 
following communications will be made : — 

Dr. J. A. Murray, F.Z.S., Acting Honorary Pathologist. 

Report on the Deaths in the Gardens during the Year 1917. 

Prof. Wood- Jones, F.Z.S., Acting Honorary Prosector. 

Exhibition of specimens illustrating the effects of Rickets. 



The following Papers have been received : — ■ 
Dr. Branislav Petronievics. 

Comparison between the Lower Jaws of the Cynodont 
Reptiles Qomphognathus and Cynognathus. 

Miss Dorothea M. A. Bate. 



On a new Genus of extinct Muscardine Rodent from the 
Balearic Islands. (Communicated by Dr. A. Smith Woodward, 
F.R.S., V.P.Z.S.) 



15 

Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary, 
Zoological Society of London, 

Regent's Park, London, N.W; 8. 
April lUh, 1918. 



No. 180 . 

ABSTRACT OF THE PROCEWDmGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON. 

April 23rd, 1918. 
A. Ezra, Esq., Vice-President, in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

Mr. D. Sbth Smith, Curator of Birds, exhibited and made 
remarks on a Zuhi Head-dress made of the plumes of the male 
Long-tailed Whydah, Chere progne. 

The Secretary (Dr. P. Chalmers Mitchell, D.Sc, LL.D., 
F.R.S.) called attention to an advertisement that had recently 
appeared in the London Press, announcing Fur Sales by Public 
Auction about to take place in the United States. The sales in 
question were only examples of what took place annually in 
London and other important commercial centres. The numbers 
advertised were smaller than usual, no doubt on account of the 
War, but they included very large quantities of animals the 
extinction of which could not be far distant, unless measures 
were carried out to protect them. In the opinion of the speaker, 
which was confirmed by the Meeting, there was urgent need for 
drastic measures to protect Mammals. The protection of Birds 
appealed to popular sentiment, and was zealously advocated by 
many influential organisations. The danger that threatened 
Mammals were even greater, and, on account of their higher 
intelligence and more sensitive nervous organisation, the cruelty 
involved in the methods of hunting, trapping, and killing them 
was incomparably greater. 



* Tliis Abstract is published by the Society at its ofEces, Zoological Gardens 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



18 

Professor Wcod-Joxes, F.Z.S., Honorary Acting Prosector, 
exhibited and made remnrks on specimens from the Prosectorium 
illustrating the eftects of Rickets. He also exhibited a set of 
anatomical prepai'ations useful for teaching purjioses, made from 
material obtained from the Society's Collection. 

Dr. A, J. Murray, F.Z.S., Acting Honorary Pathologist to 
the Society, gave a Report, illustrated by lantern-slides, of the 
causes of deaths in the Gardens during the year 1917. 

A vote of thanks was passed to Professor Wood-Jones and 
Dr. Murray for their very valuable services to the Society. 



Tlie next Meeting of the Society for Scientific Business will 
be lield on Tuesday, May 7th, 1918, at 5.30 p.m., when the 
following communications will be made: — 

E. Heron-Allen, F.L.S., F.R.M.S., F.Z.S. 

Lantern exhibition of the Arenaceous Foraminifera of the 
Genus Thurammina. 

Dr. Branislav Petronievics. 

Comparison between the Lower Jaws of the Cynodont 
Reptiles Gomphognathus and Cynognathus. 

Miss Dorothea M. A. Bate. 

On a new Genus of extinct Muscaixline Rodent from the 
Balearic Islands. (Communicated by Dr. A. Smith Woodward, 
F.R.S., V.P.Z.S.) 



List of Papers in hand : — 

C. Tate Regan, M.A.. F.Z.S., F.R.S . 

" Freshwater Fish as Food," to be taken at the Meeting on 
May 28. 



19 

Noel Taylor, B.Sc. London. 

A Oase of Hermaphroditism in a Lizard, Lacerta viridis. 
(Communicated by Prof. J. P. Hill, F.Z.S., F.R.S.) 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secreta7y. 

ZooLoaicAL Society of London, 

Regent's Park, London, JST.W. 8. 
April 30th, 1918. 



No. 181. 



I 



ABSTliA^OT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON. 

May 7th, 1918. 



Prof. E. W. MacBride, D.Sc, F.R.S., Vice-President, 
in the Chair. 



The Minutes of the last Scientific Meeting- were confirmed. 



Mr. R. I. PooocK, F.R.S., F.Z.S., Curator of Mammals, ex- 
hibited and remarked on a number of molar teeth of Elephants, 
either shed bj animals in the Society's menagerie or removed 
after death. 



Mr. E. Heron- Allen, F.L.S., F.R.M.S., F.Z.S.,gave a lantern 
exhibition of Arenaceous Foraminiferaof the genus Thurammina. 

Dr. A. Smith Woodward, F.R.S., Y.P.Z.S., in the absence of 
the authors, gave ^n account of the two following Papers : — 

D r. Branislav Petronievics . 

Comparison between the Lower Jaws of the Cynodont 
Reptiles Gomphognathus and Cynoqnatlius. (Communicated 
by C. W. Andrews, D.Sc, F.R.S., F.Z.S.) 



* This Abstract is published by the Society at its offices, Zoological Grardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings pei- annum, payable in advance. 



22 

Miss DoKOTUEA M. A. Bate . 

On a new Genus of extinct Muscardine Rodent from the 
Balearic Islands. (Communicated by Dr. A. Smith Woodward, 
F.R.S., V.P.Z.S.) 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, May 28th, 1918, at 5.30 p.m., when the 
following communications will be made : — 

Noel Taylor, B.Sc. London. 

A Case of Hermaphroditism in a Lizard, Lacerta viridis. 
(Communicated by Prof. J. P. Hill, F.Z.S., F.R.S.) 

C. Tate Regan, M.A., F.Z.S., F.R.S. 

" Freshwater Fish as Food," illustrated by lantern-slides. 



The following Paper has been received : — 
MoRLEY Roberts. 

The Function of Pathology in Evolution. (Communicated 
by the Secretary.) 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
Zoological Society op London, 

Regent's Park, London, N.W. 8. 
May UtJi, 1918. 



No. 182. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON."' 

May 28th, 1918. 



Dr. S. F. Harmer, F.R.S,, Vice-President, 
in the Chair. 



'The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a list of additions to the Menagerie in the 
month of April 1918. 

Professor J. P. Hill, F.R.S., communicated a memoir in which 
the author, Mr. Noel Taylor, B.Sc, described a case of herma- 
phroditism in the Green Lizard, Lacerla viridis. 

Mr. C. Tate Rrgan, M.A., F.E.S., F.Z.S., gave an account, 
illustrated by lantern-slides, of the Freshwater Fishes of Great 
Britain, with special reference to their value as food, and to the 
possibilities of increasing their economic use. 



* This Abstract is publislied by the Suuiety at its offices, Zoological Gardens, 
Regent's Park, N.W.. ou the Tuesday following the date of Meeting to wiiich 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of S^x Shil/higr. per annum, payable in advance. 



24 

The next Meeting of the Society for Scientific Business will be 
held on Tuesday, June 11th, 1918, at 5.30 p.m., when the 
following communications will l)e made : — 

A. Smith Woodward, LL.D., F.R.S., V.P.Z.S. 

On Two new Elnsmobi-anch Fishes from the Upper Jurassic 
Lithographic Stone of Bavaria. 

MoRLEY Roberts. 

The Function of Pathology in Ev^olution. (Communicated 
by the Secretary.) 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secrp.tary. 
Zoological Society of Loxdon, 

Regent's Park, London, N.W. 8. 
J line itL 1918. 



No. 183. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON.* 



June 11th, 1918. 



A. Ezra, Esq., F.Il.8., Vice-President, in the Chair. 



The Minutes of the last Scientific Meeting were confirmed. 

The Secretary read a list of additions to the Menagerie in the 
month of May, 1918. 

Lt.-Ool. S. MoNCKTOX OupEMAN, M.D., F.R.S., F.Z.S., described 
observations he had made in Piimrose Hill, London, N.W., on a 
colony of Burrowing Bees {Andrena fulva). 

Dr. A. Smith Woodward, LL.D., F.R.S., Y.P.Z.S., gave an 
account of his memoir entitled " On Two new Elasmobranch 
Fishes from the Upper Jurassic Lithographic Stone of Bavaria." 

Mr. MoRLEY Roberts read a paper entitled " The Function of 
Pathological States in Evolution." 



* This Abstract is published by the Society at its oiRces, Zoological Gardens. 
Regent's Park, N.W., ou the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



26 

The next Meeting of the Society for Scientific Business will 
be held on Tuesday, November 5th, 1918, at 5.30 p.m. An 
announcement of the Papers to be taken will be issued at the 
end of October. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
Zoological Society of Londox, 

Regent's Pabk, Loxdox, N.W. 8. 
June I8th, 1918. 



Exhibitions and Notices (continued). 

Mr. Or. A. BouLENGER, F.R.S. Exhibition of the head of a Congo Fisli, Hticlructjnn 

goliath 1 95 

Mr. D. SSBTH Smith, P.Z.S., Curator of Birds. Exhibition of, ;ind remarks upon, a Zulu 

Head-dress 10.") 

Dr. P. Chalmers Mitchell, F.E.S., Secretary to the Society, Remarks on an advertise- 
ment announcing Pur Sales in the United States ]95 

Professor Wood-Jones, F.Z.S., Honorary Acting Prosector. Exhibition of, and remarks 

upon, specimens from the Prosectorium illustrating the effects of Rickets 196 



PAPERS. 

1. Skull of Rana tigrina Daud. By B. L. Bhatia, M.Sc, and Baini Prashad, M.Sc. 

(Assistant Professors of Zoology, Government College, Lahore). (Text- figures 1-9.) 1 

2. Description of a new Snake of the Grenus Oligodon from Upper Burma. By G. A. 

BouLENOER, F.R.S., F.Z.S. (Text-figure 1.) 9 

3. Reptiles from the River Tajan (Trauscaspia). By L. A. Lantz. (, Plate I.) 11 

4. On the External Characters of the Lemurs and of Tardus. By R. I. Pocock, F.R.S. 

(Text-figures 1-^16.) 19 

5. A Classification of the Pyralid^iE, subfamily Hypsotropin^. Bj Sir Gkohge Hami'Son, 

Bart., F.Z.S 65 

6. First Report on the Inheritance of Visible and Invisible Characters in Silkworms. 

By Miss Mavde L. Clegiiorn, F.Z.S., F.L.S., F.E.S 133 

7. Notes on Oetacea stranded on the British Coasts during 1913-1917. By Sidney F. 

Harmer, Sc.D., F.R.S,, F.Z.S., Keeper of Zoology in the British Museum (Natural 
History) 147 

8. On the Variation of the Pit- Viper, Lachesis atrox. By Miss Joan B. Procter, F.Z.S. 

(Text-figures 1-5.) 163 

9. Report on Deaths of Animals in the Gardens in 1917. By J. A. Murray, M.D., 

B.Sc, F.Z.S., Director, Imperial Cancer Research Fund, Pathologist to the Society. . 183 



Titlepage i 

List of Council and Officers ii 

List of Contents ii 

Alphabetical List of Contributors vii 

Index XI 



PLATE. 

1918, Parts I. & II. (pp. 1-196). 



Plate 



1. 1. G7/mnodacf yius microlepis, sp.n., (^ ,'R\xerTa.JAn. 2. G. fedt- 
scJienkoi Str., (^ , Samarkand. 3. G. caspius Eichw., 9. 
JRiver Tajan 



17 



NOTICE. 



The ' Proceedings ' for the year are issued in four parts, paged consecutively j 
80 that the complete reference is now P. Z. S. 1917, p. . . . The Distribution 
is usually as follows, hut on account of war conditions Parts I. & II. are 
published simultaneously and Parts III. & IV. will probably be similarly 
issued : — 

PaS-t I. issued in March. 
,, II. ,, June. 

.. III. „ September. 

, IV. „ Dedembef. 



'Proceedings,' 1917, Parts III. & IV. (pp. 217-338), were published 
together on February 19th, 1918. 



The Abstracts of the ' Proceedings,' Nos. 175-183, are 
contained in this Parts 



PROCEEDINGS 



OF THE 



GENERAL MEETINGS FOE SCIENTIFIC BUSINESS 



OF THE 



ZOOLOGICAL SOCIETY 

OF LONDON. 



FEE is |t)2| 



iU{l ; 



\A> 



' "^S?=^t Ma3^"^ 



1918, pp. 197-310, 

WITH 2 Plates and 30 Text-figukes. 



PllINTED EOE THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT'S PAEK, 

LONDON: 
MESSRS. LONGMANS, GREEN, AND CO., 

PATERNOSTER ROW, 



LIST 

OF THE 

COUNCIL AND OFFICERS 

OF THE 

ZOOLOGICAL SOCIETY OF LONDON. 

1918. 



patron. 
His Majesty The King. 



COUNCIL. 

His Grace The Duke of Bedford, K.G., F.R.S., President 

The Hon. Cecil Baring, M.A. 
Richard H. Burne, Esq., 

M.A. 
Lt.-Col. S, Monckton Cope- 



man, M.D., F.R.S. 
Charles Drummond, Esq., 

Treasurer. 
Alfred Ezra, Esq., Vice- 
President. 
Capt. Harold S. Ferguson. 
Capt. Hugh S. Gladstone, 

M.A. 
Sidney F. Harmer, Esq., M.A., 

Sc.D., F.R.S. , Vice-President. 
Prof. James P. Hill, D.Sc, 

F.R.S. 
William Huntsman, Esq. 
Sir Edmund G. Loder, Bt,, 

Vice-Pi'esident. 



Prof. Ernest W. MacBride, 
D.Sc, LL.D., F.R.S., Vice- 
President. 

Col. Sir A. Henry McMahon, 
G.C.M.G., K.C.I.E., G.C.V.O., 
C.S.I. 

p. Chalmers Mitchell, Esq., 
C.B.E., M.A., D.Sc, LL.D., 
F.R.S., Secretary. 

Adrian D. W. Pollock, Esq. 

The Lord Queenborough. 

The Marquess of Sligo, F.S. A., 
Vice-President. 

Aubyn Trevor- Battye, Esq., 
M.A. 

Anthony H. Wingfield, Esq. 

A. Smith Woodward, Esq., 
LL.D., F.R.S., Vice-Pre- 
sident. 



PRINCIPAL OFFICERS. 

P. Chalmers Mitchell, C.B.E., M.A., D.Sc, LL.D., F.R.S., 

Secretary. 
R. I. PococK, F.R.S., F.L.S., Curator of Mammals and 

Resident Superintendent of the Gardens. 
D. Seth-Smith, Curator of Bhxls and Inspector of Works. 
Lieut. Edward G. Boulenger, Curator of Reptiles. 
Prof. H. Maxwell Lefroy, Curator of Insects. 
John Barrow, Accountant. , 
W. H. Cole, Chief Clerk. 



LIST OF CONTENTS. 

1918, pp. 197-310. 



EXHIBITIONS AND NOTICES. 

Page 
Mr. R. I. PocoCK, F.R.S., F.Z.S., Curator of Mammals. 
Exhibition of, and remarks upon, a series of the 
molar teeth of Elephants 303 

Mr. E. Heron-Allen, F.L.S., F.R.M.S., F.Z.S. Lantern- 
exhibition of Arenaceous Foraminifera 303 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of April, 1918 303 

Mr. C. Tate Regan, M.A., F.R.S., F.Z.S. Account, illus- 
trated by lantern-slides, of the Freshwater Fishes of 
Great Britain 303 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of May, 1918 304 

Lt.-Col. S.' MoNCKTON Copeman, M.D., F.R.S., F.Z.S. 
Exhibition of, and remarks upon, a colony of Burrow- 
ing Bees on Primrose Hill 304 

The Secretary. Report on the Additions to the Society's 
Menagerie during the months of June, July, August, 
and September, 1918 306 

Mr. R. I. PococK, F.R.S., F.Z.S., Ciirator of Mammals. 
Exhibition, on behalf of Mr. E. Gerrard, of the skin 
of an abnormally coloured Red Deer 307 

Prof. H. M. Lefrgy, F.Z.S., Hon. Cui-ator of Insects. 
Account, illustrated by lantern-slides, of "Wheat 
Weevil in Australia 307 



IV 

Page 
Mr. D. Seth-Smith, F.Z.S., Curator of Birds. Exhibition 

of a mounted specimen of a hybrid Cockatoo 308 

Dr. R. T. Leiper, M.D., D.Sc, F.Z.S. Lantern-exhibition 

on Diagnosis of Helminth Infections * 308 

Dr. R. T. Leiper, M.D., D.Sc, F.Z.S. Demonstration on 

the " new " Rabbit disease 308 

Prof. H. M. Lefeot, F.Z.S., Hon. Curator of Insects. 
Exhibition of a series of lantern-slides from photo- 
graphs taken in the Zoological Gardens, Sydney, 
k S. Wales 309 

The Secretary. Report on the Additions to the Society's 

Menagerie during the month of October, 1918 309 

Miss K. Lander, F.Z.S. Description and exhibition of the 

method of preparing skeletons by the use of trypsin... 310 

Mr. E. Haxschek. Description and exhibition of investi- 
gations into the forms assumed by drops and vortices 
of gelatin in various coagulants 310 

Prof. F. Wood-Jones, F.Z.S. Exhibition of a cast and set 
of Rontgen-ray photographs taken from a Chimpanzee 
which had died from pulmonary tuberculosis 310 



PAPERS. 



10. Comparison between the Lower Jaws of the Cynodont 

Reptiles Qomphognatlius and Cynognathu.s. By Dr. 
Branislav Petronievics. (Text- figures 1-8.) 197 

1 1 . On a new Genus of Extinct Muscardine Rodent from 

the Balearic Islands. By Dohothea M. A. Bate, 
Hon. M.B.O.U. (Plate I. and Text-figures 1 & 2.)... 209 

12. A Case of Hermaphroditism in a Lizard, Lacerta 

viridis. By JSToel Tayler, B.Sc. (Lond.). (From 
the Zoological Department, University of London, 
University College.) (Text-figures 1-3.) 223 



Page 

13. On two new Elasniobranch Fishes {Crossorhinus juras- 

sicus, sp. nov., and Protospiiiax annectaii,s, gen. et sp. 
nov.) from the Upper Jurassic Lithographic Stone 
of Bavaria. By Arthur S ith Woodward, LL.D., 
F.R.S., V.P.Z.S. (Plate I.) 231 

14. The Function of Pathological States in Evolution. By 

MoRLEY Roberts 237 

15. Notes on the Beavers at Leonardslee, 1916-1918. By 

Sir Edmund G. Loder, Bart., Vice-President Z.S. ... 255 

16. On the Madagascar Frogs of the Genus Mantidactyhis 

Blgr. By G. A. BouLENGER, F.R.S., F.Z.S '..... 257 

17. Ciliary Action in the Internal Cavities of the 

Ctenophore Flerr.robrachia pileus Fabr. By James 
F. Gemmill, M.A., M.D., D.Sc, F.Z.S. (Text- 
figures 1 & 2.) 263 

18. On Seymouria, the most primitive known reptile. By 

D. M. S. Watson, M.Sc, Capt. R.A.F., Lecturer 
in Vertebrate Palaeontology, University College, 
London. (Text-figures 1-15.) 267 

Alphabetical List of Contributors vii 

Index .^ , xi 



ALPHABETICAL LIST 



CONTRIBUTORS, 

With References to the several Articles contributed hy each. 
(1918, pp. 197-310.) 



Page 

Bate, Miss Dorothea M. A., Hon. M.B.O.U. 

On a new Genus of Extinct Muscardine Rodent from 
the Balearic Islands. (Plate I. and Text-figures 1 & 2.)... 209 

BouLENGER, G. A., F.R.S., F.Z.S. 

On the Madagascar Frogs of the Genus Mantidactylus 
Blgr. 257 

COPEMAN, Lt.-Col. S. MONCKTON, M.D., F.R.S., F.Z.S. 

Exhibition of, and remarks upon, a colony of Burrow- 
ing Bees on Primrose Hill 304 

Gemmill, James F., M.A., M.D., D.Sc, F.Z.S. 

Ciliary Action in the Internal Cavities of the Cteno- 
■^\i.ovQ Pleurohrachia pileus 'PaJoY. (Text-figures 1 & 2,)... 263 

Hatschek, E. 

Description and exhibition of investigations into the 
forms assumed by drops and vortices of gelatin in various 
coagulants 310 

Heron- Allen, E., F.L.S., F.R.M.S., F.Z.S. 

Lantern- exhibition of Arenaceous For^minifera 303 



Vlll 

Page 
Lander, Miss K., F.Z.S. 

Description and exhil)ition of the method of preparing 
skeletons by the use of trypsin 310 

Leproy, Prof. H. M., F.Z.S. , Hon. Curator of Insects. 

Account, illustrated by lantern-slides, of Wheat Weevil 
in Australia 307 

Exhibition of a series of lantern-slides from photo- 
graphs taken in the Zoological Gardens, Sydney, N. S. 
Wales 309 

Leiper, Dr. R. T., F.Z.S. 

Lantern-exhibition on Diagnosis of Helminth Infec- 
tions 308 

Demonstrations on the "new" Rabbit disease 308 

LoDER, Sir Edmund G., Bai-t., Vice-President Z.S. 

Notes on the Beavers at Leonardslee, 1916-1918 255 

Mitchell, P. Chalmers, C.B.E., M.A., LL.D., F.R.S., 
F.Z.S., Secretary to the Society. 
Report on the Additions to the Society's Menagerie 
during the month of April, 1918 303 

Report on the Additions to the Society's Menagerie 
during the month of May, 1918 304 

Report on the Additions to the Society's Menagerie 
during the months of June, July, August, and September, 
1918 306 

Report on the Additions to the Society's Menagerie 
during the month of October, 1918 309 

Petronievics, Dr. Branislav. 

Comparison between the Lower Jaws of the Cyno- 
dont Reptiles Gomphognathus and Gynognathtts. (Text- 
figures 1-8.) .' 197 



IX 

Page 
PocooK, R. I., F.R.S., F.Z.S., Curator of Mammals. 

Exhibition of a series of the molar teeth of Elephants. 303 
Exhibition, on behalf of Mr. E, Gerrard, of the skin 
of an abnormally coloured Red Deer 307 

Regan, C. Tate, M.A., P.R.S., F.Z.S. 

Account, illusti-ated. by lantern-slides, of the Fresh- 
water Fishes of Great Britain 303 

ROBEKTS, MOELEY. 

The Function of Pathological States in Evolutioiu 237 

Seth-Smith, D., F.Z.S., Curator of Birds. 

Exhibition of a mounted specimen of a hybrid 
Cockatoo 308 

Tayler, Noel, B.Sc. (Lond.). 

A Case of Hermaphroditism in a Lizai'd, Lacerta 
viridis. (Text-figures 1-3.) 223 

Watson, D. M. S^ M.Sc, Capt. R.A.F. 

On Heymouria, the most primitive known reptile. 
(Text-figures 1-15.) 267 

Wood- Jones, Prof. F., F Z.S. 

Exhibition of a cast and set of Rontgen-i-ay photo- 
graphs taken from a Chimpanzee which had died from 
pulmonary tuberculosis 310 

Woodward, Dr. A. Smith, F.R.S., V.P.Z.S. 

On two new Elasmobranch Fishes {Crossorhimis juras- 
sicus, sp. nov., and Frotospinax annectans, gen. et sp. no v.) 
from the Upper Jura-ssic Lithograpliic Stone of Bavaria. 
(Plate I.) 231 



Proc. Zool. Soc. — 1918. 



INDEX. 



1918.— Pages 197-310. 



[New names in clarendon type. Systematic references in italics 
(z. s.L.) indicates additions to the Society's Menagerie.] 



Alligator mississippiensis (z. s. l.), 309. 
Ammotragus lervia (z. s. l.), 303. 
Andrena fulva, 304. 
Ates: 

Exhibition of a mounted specimen of 
a hybrid Cockatoo, 308. 

Calliope calliope (z, s. i:.\ 304. 
Capra segagrus (z. s. l.), 304. 
Cercopithecus ascaniiis (z. s. l.), 309. 
Chamteleon calcaratus (z. s. l.), 304. 
Connochfetes albojubatus (z. s. l.), 306. 
Crossorhinus jurassicvis, sp. n., 
231. 

Equus kiang (z. s. l.), 307. 

Gecco verticillatus (z. s. l.), 307. 

GEOGRAnilCAL: 

Madagascar Frogs of the genus Man- 
tidactyliis, 'Ibl. 

Hemitragus jemlaicus (z. s. L.), 307. 
Hypnomys, gen. n., 210. 

mahonensis, sp. n., 218. 

morpheus, .sp. n., 219. 

Hystrix cristatus (z. s. l.), 3u7. 



Insecta : 

Observations on a colony of burrowing 
Bees, exhibited, 304 ; On Wheat 
Weevil in Australia, 307. 

Lacerta taurica (z. s. l.), 306. 

Mammalia : 

On a new genus of Extinct Muscar- 
dine Rodent from the Balearic 
Islands : structure, systematic, 
209 ; Notes on the Beavers at 
Leouardslee, 255 ; The ages of 
Elephants, as inferred from the 
Mular Teeth, exhibited, 303; An 
abnormally coloured Red Deer, 
skin exhibited, 307; The "New" 
Rabbit Disease, demonstration, 
308. 
Mantidactylus ambohimitcmbi, 

sp. n., 260. 

MOLLUSCA ; 

Ciliary action in Pleurobrachia pileus, 
structure, 263 ; Lantern-exliibition 
of Ai-enaceous Foramii.ifera, 303. 

Morphology. See Structure. 

Myocastor coypus (z. s. L.j, 307. 



X}1 



INDEX. 



Pal;\>ornis cyanoecpliala (z. s. l.), 303, 

Pathology. 

The function oC Pathological stales 
in Evolution, 237; Diagnosis of 
Helminth infections, 308. 

Pisces : 

On two new Elasmobranch Fishes : 
systematic, 231 ; Account of the 
Freshwater Fishes of Gi-eat 
Britain, 303. 

Protospinacidge, fam. n., 232. 

Protospinax, gen. n., 233. 

annectans, sp. n., 233. 

Python reticuhxtus (z. s, i..), 309. 



Eeptilia : 

Comparison between the lower jaws 
of the Cynoclont Reptiles Gomjjho- 
gnathus and Cipiognaihus: struc- 
ture, 197; On Hermaphroditism in 
a Lizard : structure, 223 ; Mada- 
gascar Frogs of the genus Manti- 
dactylus : structure, systematic. 



257 ; On Sci/mouria, the most 
primitive known reptile : structure, 
267. 



Structure. 

Mammalia: On a new genus of ex- 
tinct Muscardine Rodent from the 
Balearic Islands, 209. 
Rei'tilta : Comparison between the 
lower jaws of the Cynodont Rep- 
tiles Gompliognafhus and Cyno- 
gnathus, 197 ; On Hermaphro- 
ditism in a Lizard, 223; Mada- 
gascar Frogs of the genus Manti- 
dacfylus, systematic, 267 ; On 
8eymouria, the most ])rimitive 
known reptile: structure, 267. 
MoLLUSCA : Ciliary action in Pleuro- 
brachia pileus, 263. 



Taurotragus oryx (z. s. l.), 307. 



PRINTED IIY TAYI.0E AND FRANCIS, RED l.:0N COURT, FLEET STRKET. 



L«==^— =— — ff 



PROCEEDINGS 

OF THE 

OENERAL MEETINaS EOR SCIENTIFIC BUSINESS 

OF THE 

ZOOLOGICAL SOCIETY 

OF LONDON. 
1918. 



PARTS III. & IV. 



CONTAINING PaGES 197 TO 310, WITH 2 PlATES 

AND 30 Text-figures. 

r 




PRINTED FOR THE SOClET^j iv-i^ae^-" 
SOLD AT ITS HOUSE IN EEGENT'S PARK. 

LONDON : 
MESSRS. LONGMANS, GREEN, AND CO., 

PATERNOSTER ROW. 



[Price Twelve Shillings. 



LIST OF CONTENTS. 
1918, Parts III. & IV. (pp. 197-310). 



EXHIBITIONS AND NOTICES. 

Mr. R. I. PococK, F.R.S., F.Z.S., Curator of Mammals. Exhibition of, and remarks upon, 

a series of the molar teeth of Elephants 303 

Mr. E. Hkron-Allen, F.L.S., F.R.M.S., F.Z.S. Lantern-exhibition of Arenaceous 

Foraminifera 303 

The Secketary. Eeport on the Additions to the Society's Menagerie during the month 

of April, 1918 303 

Mr. C. Tate Eegan, M.A., F.E.S., F.Z.S. Account, illustrated by lantern-slides, of the 

Freshwater Fishes of Great Britain 303 

The Secretary, Eepoi-t on the Additions to the Society's Menagerie during the month 

of May, 1918 304 

Lt.-Col. S. MoNCKTON CopEMAN, M.D., F.E.S., F.Z.S. Exhibition of, and remarks upon, a 

colony of Burrowing Bees on Primrose Hill 304 

The Secretary. Eeport on the Additions to the Society's Menagerie during the months 

of June, July, August, and Septenfber, 1918 306 

Mr. E. I. PococK, F.E.S., F.Z.S., Curator of Mammals. Exhibition, on behalf of Mr. E. 

Gerrard, of the skin of an abnormally coloured Eed Deer 307 

Prof. H. M. Lefroy, F.Z.S., Hon. Curator of Insects. Account, illustrated by lantern- 
slides, of Wheat Weevil in Australia 307 

Mr, D. Seth Smith, F.Z.S., Curator of Birds. Exhibition of a mounted specimen of a 

hybrid Cockatoo 308 

Dr. E. T. Leiper, M.D., D.Sc, F.Z.S. Lantern- exhibition on Diagnosis of Helminth 

Infections 308 

Dr. E. T. Leiper, M.D., D.Sc, F.Z.S. Demonstration on the " new " Eabbit disease 308 

Prof. H. M. Lefroy, F.Z.S., Hon. Curator of Insects. Exhibition of a series of lantern- 
slides from photographs taken in the Zoological Gardens, Sydney, N. S. Wales .... 309 

- Contents continued on page 3 of Wrapper. 



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FEB 17 1920 

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ZOOLOGICAL SOCIETY OF LONDON. 



PAPERS. 



10. Comparison between the Lower Jaws of the Cjnodont 
Ee})tile3 Gompkog^iatlms and Cjjnoi/nathus. By Dr. 
Beanislav Petkonievics *. 

[Received March 6, 1918 : Read May 7, 1918.] 
(Text-figures 1-8.) 

Being occupied at the end of last year (1917), when in London, 
with the problem of the double articulation of the lower jaw, 
which is to be expected in the direct reptilian ancestors of the 
Mammals, I was led to examine the lower jaws in the specimeKs 
of the Theriodont Reptiles that are preserved in the Natural 
Histoi-y Museum. One of the skulls in question, that described 
by Seeley in 1895 as one of the two specimens of GomphognathKs 
from Lady Frere (S. Africa), struck me on account of the strange 
appearance of its lower jaw. On comparing carefully this latter 
with the lower jaw in Cynognatlius a-ateronotiis, described by 
Seeley in 1895, 1 have arrived at the conclusion that the difference 
between them is a very considerable one, and that in the lower 
jaw of Goviphognathus we have the greatest known development 
of the dentary bone in any theriodont reptile, a development 
representing the initial state of the double articulation, if not the 
actual double articulation itself. 

The main points of diflterence in the lower jaws of Gompho- 
gnathus and Cynognathus are the following : — 

1. In Gomjjhognathus the articular bones are placed inside and 
laterally at some distance from the dentary bone, whilst in 

* Comramiicated by Dr. C. W. AndeewSj F.R.S., F.Z.S. 

Proc. Zool. Soc— 1918, No. XV. 15 



198 



DR. B. PETRONIEVICS ON THE LOWER JAWS OF TUE 



Cynognathus they are attached closely to the dentaiy bone 
(coinp. the riglit ramus of the lower jaw of Gomj^Jtoynailivs 
in text-fig. 1 with the right ramus of Cynognathus in text- fig. 7). 



Text-figure 1. 



art.s 
arl.srf.dent.?! 



dentri 




Gomphognatlms : inferior surface of skull and mandibles. X i- 

ang. ; angular. 

ang.lm. ; angular lamina. 

ang.n. ; angular notch. 

art. ; articular. 

art.srf.dent. ? ; articular surface of dentar3r. 

art.srf.qu. ; artieular surface of quadrate. 

dent. ; dentary. 

dent.n. ; dental notcli. 

pra>'t. ; prearticular. 

sang. ; supra- angular. 

spl.; splenial. 

St. ? ; stapes. 

2. In Gomphognathus the posterior end of the dentary hone 
goes almost as far back as the articidar bones (conip, text-figs. 2 



CYNODONT REPTILES GOMPHOOxNATnUS AND CYNOGNATHUS. 199 



and 4), whilst in Cynognathus the articular goes farther back 
than the clentary * (and it does this more externally than internally 
— couip. text-fig. 5 and text-fig. 6, in which the length of these 
portions of the dentary bone is the same). 

3. In Gomphognathus the posterior edge of the dentary bone 
covers externally the articular bones almost entirely (comp. text- 
figs. 2 and 4), whilst in Cynognathibs the hind part of the articular 
bones is clearly visible and considerably external to the dentary 
bone (text-tig. 5). 

Text-figure 2. 



pre on 



in. con 




pr.ang. 



Gompliognatlms : right ramus of lower jaw, external view. X i- 
Shaded part covered with matrix. 

dent. ; dentary. 

iii.con.dent. ; incipient condj'le of dentary. 
pr.cor. ; coronoid process of dentary. 
pr.ang. ; processus angularis. 

In connexion with the first point we must add that Seeley's 
contention (comp. Seeley(l), 1895, p. 26), that the articular bones 
of GompJwgnathus were "somewhat displaced, being drawn away 
laterally from the edge of the dentaiy bones," must be wholly 
rejected, because the lateral distance of these bones from the 
dentaries is exactly the same on both sides (comp. text-tig. 1), 
and the bony connexion of the almost hoiizontally situated 
reflected angular lamina, which is well preserved on the left side, 
is such (comp. text-fig. 1) that no doubt about the naturalness of 
its position can be entertained. The angular notch limited by 
this lamina is a large one, and comparatively large, also, is the 
notch lying iu front of it, between the dentary on the one side, 
and angular and splenial on the other (comp. text-fig. 1 ; the hind 
part of the splenial is preserved only on the right side). 
In Cynognathus the reflected angular lamina, whose j30sition is 
almost a vertical one, is preserved only partially (comp. text-fig. 6 

•* In Cynognathus platyceps the hind end of the dentary is more prolonged back- 
wards than in Cynognathus crateronottts, but it did not reach the squamosal. 
Comp. Seeley (2)/l895, fig. 29, p. 135, and Eroom, 1904, text-fig. 100, p. 496. 

15* 



200 



DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE 



and text-fig. 7), but it must have been relatively smaller than 
in Gomphogyiathus (comp. text-fig. 6). 

In Gomphoynathus there is a bony connexion between the 
articular bones and the dentary (whose thickness could not be 
established, but which probably is not a coDsiderable one), this 
bony connexion, forming the rodf of the angular notch, being 
well preserved and visible on the left side (comp. text-fig. 1). The 

Text-figure 3, 



m. con. 




Gom/pliognathus : right ramus of lower jaw, internal view. X J. 
Shaded part covered with matrix. 

ang. ; angular. 

art. ; articular. 

cr. ? ; coronoid. 

dent. ; dentary. 

in.con.dent.; incipient condyle of dentary. 

jirart. ; prearticular. 

spl. ; splenial. 



Text-figfure 4. 




in.condent 



ang. 



jjr.ang. 



Gompkoffnatlnts : left ramus of lower jaw, external view. X j. 

ang. ; angular. 

dent. ; denrary. 

in.con.dent. ; incipient condyle of dentary. 

pr.ang. ; processus angularis. 

sang. ; supra-angular. 



sutures between the articular bones are not to be traced in their 
whole length in Go^njyhoguathits (comp. text-figs. 3 and 4;, while 
they are clearly visible in Gynognathus (comp. text-figs. 5, 6, 



CYNODONT REPTILES GOMPHOaNATHUS AND CYNOGNATHUS, 201 

and 7) : it is probable that the mutual limits of these bones are 
in both cases the same. The suture between the articular and 
prearticular is obliterated in both cases. 

As to the second point. The coronoid process of the left 
dentary in Gomphognathus is only partially preserved (text-fig. 4), 

Text-fiefure 5. 




Cynognatlius oiyiteronotws i right ramus of lower jaw, external view. 
Shaded part covered with matrix. 

anff.\ ano:ular. 
art. ; articular. 
dent. ; dentary. 
sang. ; supra-angular. 



Text-ficfure 6, 



dent. 




ang. , ang.TL. 

ana. Jrn 

Cynognatlius craieranotus : right ramus of lower jaw, internal view. X 
Shaded part covered with matrix. 



anff. ; angular. 
ang.lm. ; angular lamina. 
ang.n. ; angular notch. 
art. ; articular. 
cr. ; coronoid. 



dent. ; dentary, 
prart. ; pre-articular. 
sang. ; surangular. 
spl. : splenial. 



whilst the right .dentary shows this process in its entirety, and 
although the upper part of it is covered with matrix, its upper 
edge is clearly visible from above (com p. text-fig. 2). As the hind 



202 



DR. B. PETROXIEVICS ON THE LOWER JAWS OF THE 



end of tliis eilge lies in direct prolongation of the lower part of 
the posterior edge of the coronoid process uncovered by the new 





Cynognathus crateronotus : right ramus of lower jaw, 
inferior surface. X f. 



ang. ; angular. 
ang.lm. ; angular lamina, 
ang.n. ; angular notcli. 
art. ; articular. 



dent. ; clentary. 
prart. ; pre-articular. 
sang. ; suraugular. 



preparation, so the naturalness of this posterior edge and conse- 
quently of the prominent backward condylar prolongation of the 
right dentary (text-fig. 2) cannot possibly be doubted. As this 



CYNODONT REPTILES GOMPnOGNATHUS AND CYNOGNATHUS. 203 

condylar prolong.ation is damaged beneath and at its posterior end, 
whose transverse section shows a triangular shape, the articular 
surface of the condyle cannot be certainly established. On the 
left side (text-fig. 4) only a part of the prominent backwai-d condylar 
prolongation is preserved, so that on the outer side of the left 
jaw the hind end of the articular bones seems to go farther back 
than the hind end of the dentary, while on the right side the 
reverse seems to take place. But the hind end of the articular 
bones on the right side is evidently damaged, so that it is not 
impossible that the dentary reached As far back as the articular 
bones, and that a double articulation really did take place in 
Gomphog7iathus*. 

The possibility of the double articulation in Gomphognaihus 
follows also from the fact of two different articular surfaces 
preserved in our specimen on the left side (comp. text-fig. 8). 
A comparison with the corresponding region in the type skull of 
Diademodon described by Watson (comp. fig. 3 in Watson 1911) 
shows (comp. also fig. 8 in Watson 1911 representing the quad- 
ra'te in Goinphognathus jr,olyp]iagus) that only the triangular 
hollow on the front face of the squamosal, in which the missing 
quadrate was received, is preserved in our specimen on the left 
side together with the two notches which received the two pro- 
cesses of the quadi-ate (these notches being filled with matrix). 
As there is in our specimen on the left of these notches and 
in a more forward position another flat surface on the squa- 
mosal, which lies in the same direction as the hind end of 
the dentary (comp. text-figs. 1 and 8), it is quite possible that 
into this flat surface was received the hind end of the condylar 
prolongation of the dentary. Indeed, as the articular end in 
our specimen did not reach the squa,mosal on the external side 
of the dentary as in Gomphognathus kannevieyeri (comp. fig. 1 in 
Seeley (1) 1895, p. 5, and especially fig. 2 in Broom, 1904, 
pi. xxxv.), or as in Cynognathus (comp. fig. 8 in Seeley (2) 1895, 
p. 81, and fig. 1 in Broom, 1904, pi. xxxv.), so the flat surface in 
question could be reached in our specimen only by the hind end 
of the dentary (supposing that it was reached by the lower jaw 
at all). 

The third point is a corolloTy of the second. As the posterior 
edge of the left dentary of GompJwgnathus, extending from the 
angle up to the condylar prolongation, is almost undamaged 
(comp. text-fig. 4), whilst the corresponding posterior edge of the 
right dentary is not inconsiderably damaged, the difference in the 
extent of the covering of the articular bones by the dentary from 
outside in Gomphognathus and Cynognathus is strikingly shown 

* The length of the two dcntaries of Gomjiho/fnafhus, as they are preserved, is 
almost the same (149 mm.), when measured from the hind point of their symphj'sis. 
But a certain asymmetry of them is not iniprohahle, hrcause when we compare the 
length of the upper edge of the left dentary wiih the corres])onding line in the 
coronoid process of the right deutar3-, we tind the first to he 63 and the second 
66 mm; 



204 DR. B. rKTRONIEVICS ON THE LOWER JAWS OF THE 

when text-fig. 4 is com pared witli text-fig. 5. So that, altiiough the 
]iind end of the dentnry in Cynognathns goes farther back on tlie 
inner tlian on the outer side, it is clear from this comparison that it 
did not reach tlie squamosal and that the possibility of an incipient 
comlyle of the dentary in Cynognathits is excluded. 

Text-figure 8. 

arLsfdent ^ 

I 

I QJt.srt 




dent. 



G-ow plwjna tints : avticular surface of the left squamosal. Nat. size. 

art. ; avticnlar. 

art.sf.dent ?; probable articvilav sui'face of dentary. 
art.srfqii. ; articular surface of left quadrate. 
dent. ; dentary. 

The inner side of the jaws shows the same bones in both 
cnses (comp. text-tig. 3 with text-fig. 6). The comparison shows 
that the splenial goes farther back in Gomphognathus than in 
Cynognathiis. 

The essential difference above mentioned that exists in respect 
to the relation of the articular bones and the hind end of tlie 
dentary to the squamosal between the Gomphognathiis-^VvM of our 
specimen a,nd the other specimens known under the names of 
GomphognatliKS and Biademodon (a general survey of which has 
been given by Watson, 1911, p. 327 seq.) suggests the well- 
founded supposition that our specimen is generically different 
from all the other specimens of the same kind, which are all 
essentially similar in the above point. Beyond that, a comparison 
between the lower jaw of Gomphognathus kannemeyeri (figured 
by Seeley (1) 1895, fig. 2, p. 8) and of Diademodon mastacus 
(fignred by Broom, 1905, fig. 1, pi. x.) shows that they are similar 
in shape, while the shape of the lower jaw of our sjjecimen 
(comp. text-fig. 2), especially the shape of its coronoid process, is very 
different. So that I agree with Watson in his identification of 
Gomphogaatlms and Diademodon for all the other specimens 



CYNODONT REPTILES QOMPHOGNATHUS AND CYNOGNATHUS. 205 

except the one in question, which alone has to retain the old 
geneiic name of Gomphognathiis* . 

Having tinished the comparison between the lower jaws of 
Gomphognathus and Cynognathus, I will add here some general 
remarks concerning the origin of the mandibular articulation and 
the origin of the Ossicula auditus in Mammals, inasmuch as these 
two veiy closely connected problems receive new light from oar 
comparison. 

That the mandibular articulation in the most primitive mammals 
(the hypotheticid order Promammalia of Gregory — comp. its 
definition in Gregory, 1908, p. 164) was a double one, our 
Gomphognathus-iikuW puts this point beyond question, as it shows 
that in this advanced Theriodont the dentary almost if not 
actually reached the squamosal. That the dentary articulation 
was situated in the same plane with the articular articulation, and 
vot in front, is also a point put beyond question. That the two 
articulations have worked closely together and practical 1}' as one 
articulation, is also very probable (on these three points in 
primitive Mammals comp. Gregor}"^, 1908, pp. 135-138). 

Our Gom])hognathus-'s^\\\\ shows also the way in which the 
cotylus and condylus of the dentary articulation in Mammals 
have probably arisen. As the articular surface of the dentary 
situated on the squamosal is a smooth plane surface in our 
specimen, so we have to suppose that also the corresponding 
surface on the condylar prolongation of the dentary was a smooth 
plane surface. Consequently we may suppose that this latter 
surface was a mechanical lesult of the latei-al movements of the 
lower jaw, through which the hind end of the dentary {%. e. the 
condylar prolongation of its ascending process) was brought in 
contact with the squamosal. The primitive condition of the 
dentary articulation in Mammals would, according to this 
supposition, be a simple syndesmosis, as we find svich a syndes- 
mosis secondarily in Tatusia hyhricla among living forms. The 
different forms of the mandibular articulation in higher Mammals 
would then be considered as further mechanical results of jaw- 
movements according to the mechanical theory of normal articu- 
lations of Fick t, Tornier j, and Cope §. 

Still greater is the importance of our specimen in respect to 

* Between the skulls of Cynognathus on the one side and the skulls of Diade- 
modon (and perhaps also of Gomphognathus) on the other, 1 find an essential 
difference in a separate ossification that exists in front of the epipter3'goid and below 
the postfrontal bone in the British Museum specimen R 3587 of Diademodon- 
skull (described by Watson, 1911, who has not recognized its separate nature), an 
ossification that 1 suppose to be an incipient orbitosphenoid. The front edge of 
the epipterygoid bone in Cynognathus crateronotus is damaged, but the under 
surface of the postfrontal bone is so smooth, that a correspondmg separate orbito- 
sphenoidal ossification is a very improbable one (comp. fig. 6 in Watson, 1911, p. 300, 
with fig. 6, p. 76, in Seeley (2), 1895). 

f Comp. R. Fick, " Ueber die Form der Gelenkflachen " in ' Arcliiv fiir Anatomie 
und Physiologic,' 1890, p. 391. 

X Comp. J. Tornier, "Das Entstehen der Gelenkformen " in ' Archiv fiir Ent- 
wickelungsniechanik,' 1897, pp. 124-158, 224-268, 307-346. 

§ Comp. E. Cope, 'Primary Factors of Organic Evolution,' 1904, p. 283 seq. 



206 DR. B. PETRONIEA'ICS ON TFIE LOWKIl JAWS OF THE 

the qiiestion of the origin of tlie Ossicula auditns in Mammals. 
Our specimen shows the beginning of the separation of the 
articular bones from the dentary, a state of things that the 
classical theory of the origin of Mammalian ossicula auditus 
necessarily presupposes. As we have seen, the bony connexion 
between the articular bones and the dentary is a r-elatively much 
reduced one, especially in the hind part (comp. text-fig. 1). The fact 
that the quadrate has been lost in our specimen on the left siile, 
where it might have been preserved, shows that it was probably 
only insulticiently fixed in the corresponding groove of the 
squamosal. The movable condition of the quadrate, that the 
classical theory presupposes, is here, as it seems, also in its 
beginning. And finally we find in our specimen on the left side 
a displaced bone (comp. text-fig. 1, st.), that is, according to 
another specimen, to be considered as the bone connecting the 
region of the fenestra vestibuli with the quadrate, consequently 
as the stapes or the columella (on this bone comp. Seeley (1 
1895, p. 25, Broom 1904, p. 49], who considers it in Ci/noc/nathus 
platyceps wrongly as a tympanic ring, and especially Watson, 
1911, p. 324). 

From the above it follows that we have in Gomphogiuithns 
realized all the elements that constitute the morphologically 
initial state in the evolution of the mammalian ossicula auditus, 
and we have only to suppose that in the descendants of the cor- 
resiponding relatives of Gomphognathus (the direct ancestors of 
Mammals) this state of things increased in the direction of a 
furtlier separation of the stapes-quadrate-articular bone chain 
from the squamosal and dentary bones, in order to reach ,the final 
state, when in true Mammals the former were transformed into 
the ossicula auditus. Into the question how this ti^ansformation 
took p)lace, and into the corresponding question of the homologies, 
I cannot enter here (comp. the Literature, Nos. 7-11). I mention 
only that the tympanic membrane of the Mammals is most pro- 
bably a neomorph (and I agree in this point with Gaupp — comp. 
Gaupp, 1911, p. 641 seq. and p. 659, while in many other con- 
clusions readied by Gaupp — comp. p. 633 and p. 656 — I must 
disagree), and that the manubrium mallei in Mammals cannot be 
homologized with the extra-col umella of the Reptiles, as Kings] ey 
does (comp. Kingsley, 1900, p. 232 seq.). 

In finishing this paper I desire to express my thanks to 
Dr. Smith Woodward and Dr. Andrews, of the British Museum 
(Natural History), for the loan of t\\e new preparations (executed 
by Mr. Hall), and to Dr. Andrews for some valuable help. 

Literature. 

1. H. G. Seeley. — Researches on the Structure, Organization, 
and Classification of the Fossil Reptilia. Part IX. Section 4. 
On the Gomphodontia. In ' Philosophical Transactions 
Royal Society,' 1895, vol. 186, B, pp. 1-58 (On Gompho- 
gnath us, pp. 3-31). 



CYNODONT REPTILES GOMPHOGNATHUS AND CYNOG>f ATHUS. 207 

2. H. CI. Seeley. — Researches, etc., Part IX. Section 5. On 

the Skeleton in new Ojnodontia from the Karoo Rocks. In 
' Philosophical Transactions Royal Society,' 1895, vol. 186, B, 
pp. 59-148. 

3. R. BaooM. — On the Structure of the Theriodont Mandible 

and on its Mode of Articulation with the Skull. In ' Pro- 
ceedings of the Zoological Society of London,' 1904, vol. i., 
pp. 490-498. 

4. R. Broom. — On some Points in the Anatomy of the Theriodont 

Reptile Diademodon. In ' Proceedings of the Zoological 
Society of London,' 1905, vol. i., pp. 96-102. 

5. D. M. S. Watson. — The Skull of Diademodon, with Notes on 

those of some other Oynodonts. In ' Annals and Magazine 
of Natural History,' vol. viii., 1911, pp. 293-330. 

6. D. M. S. AVatson. — On some Reptilian Lower Jaws. In 

'Annals and Magazine of Natural History,' vol. x., 1912, 
pp. 573-587. 

7. R. Broom. — On the Fate of the Quadrate in Mammals. In 

' Annals and Magazine of Natural History,' vol. vi., 1890, 
pp. 409-411. 

8. J. S. Kingsley. — The Ossicula auditus. In ' Tufts College 

Studies,' No. 6, 1900, pp. 203-274. 

9. H. Gadow — The Evolution of the Auditory Ossicles. In 

' Anatomischer Anzeiger,' Bd. xix., 1901, pp. 396-411. 

10. W. K. Gregory. — The Orders of Mammals. In ' Bulletin of 

the American Museum of Natural History,' vol. xxvii., 
1910, chp. I : "The Origin of Mammals and the Problem 
of the Ossicula auditus," pp. 113-143 (comp. also Literature 
on " Evolution of the Mammalian Ossicula auditus," 
pp. 484-5). 

11. G. E. Gaupp. — Beitrage zur Kenntniss des Unterkiefers der 

Wii"beltiere. III. Das Problem der Entstehung eines 
" secundiiren " Kiefergelenkes bei den Saugern. In 
'Anatomischer Anzeiger,' Bd. xxxix,, 1911, pp. 609-666. 



p. Z. S. 1918, Miss BATE, PI. I. 







4.x 4- 





II. ' ! 1x4 
6.x ^ 



Z x3 




5.X4. 





\ 
3x4 



v._:' 





9x4 




12. I f^4 |5.\f ix4 
7x3 



^-^' 

"^ 



/. .1/. JVoodzvard, ,h'i „,„ 

HYPNOMYS, gen. nov. et LEITHIA. 




^ \ 



14x2. 



S nanielsson, Ltd. 



A NEW GENUS OF EXTINCT MUSCARDINE RODENT. 209 

11. On a new Genus oE Extinct Muscardine Rodent from 
the Balearic Islands. By Dorothea M. A. Bate *, 
Hon. M.B.O.U. 

1 Received April 9, 1918 ; Read May 7, 1918.] 
(Plate I.t and Text-figs. 1,2.) 

Ikdbx. 

Systematic .- Pare 

Hypnomys, gen. n 210 

H. mahonensis, sp. ii 218 

S. morpheus, sp. n 219 

During a second visit to Mallorca in 1 910 in search of ossiferous 
deposits a few rodent remains were obtained ; the following year 
further researches were carried out in Menorca by means of a 
grant from the Trustees of the Percy Sladen Memorial Fund, iind 
here similar remains were found to be somewhat more plentiful, 
occurring in several fissures in the Miocene limestone. A de- 
scription of the deposits from which the collection was obtained 
has already been published %. A first cursory examination 
of the specimens led me to suppose that they represented large 
forms of Uliomys§ or Leitkia\\. Since then a number of spe- 
cimens from Menorca have been developed from the hard matrix 
in which they were embedded, and all have been carefully 
examined, with the result that it is found that they cannot be 
included in any genus with which I have been able to compare 
them. The examples from the two islands diff"er considerably in 
size, those from Menorca being the larger, and they are probably 
specifically distinct: in this connection it is interesting to remember 
that, after the examination of a very large quantity of material, a 
similar variation in size was found to obtain in Myotragi(,s %. 
This seems to point to a longer period of isolation in Menorca, 
the most easterly and probably the first of the group to be 
separated from the mainland. 

All the remains obtained are now in the collection of the 
British Museum (Natural History). No complete skulls and 
only a few limb-bones were procured, but it will be seen from 
the descriptions of the specimens given below that the Balearic 
genus should undoubtedly be included in the Muscardinidse. 
The specimens to be described are intermediate in size be- 
tween the largest recent forms and the extinct Leithia from 
Malta, and show a number of points of resemblance both to 
the recent Eliomys and to the Maltese Leithia, but at the same 
time difiier to an equal extent from both these genera. The 

* Communicated by Dr. A. Smith Woodwahd, F.R.S., F.Z.S. 
+ For explanation of the Plate see p. 222. ^ 

I Geol. Mas:. [6] vol. i. 1914, pp. 337-45. 

§ ihu. p. ibo. 

II Proc.Zool.Soe. 1916. p. 424. 

•j[ See Andrews, Phil. Trans. Roy. Soc. ser. B, vol. 206, 1915, p. 801. 



210 MISS DOROXnEA BATE ON A KEW GENUS 

range of species in neighbouring islands is always of interest, 
whicli may perhaps make it worth mentioning here the present- 
day distribution of Eliomys in the Balearics. Mr. Oldfiekl 
Thomas has described E. gymnesicus* from Menorca, a species 
smaller than, though similar in coloration to, E. quercinus^ which, 
in spite of this, he believes to be most neai'ly allied to the Southern 
forms. In an earlier paper t the same author mentions that this 
animal is also well known in Mallorca but is said not to occur in 
Ibiza, Unfortunately I have not seen one from the largej- island, 
but in the British Museum there are now several specimens from 
the small island of Formentera to the south-west of Ibiza, which 
prove to belong to the large form E. lusitanicus which is found 
on the neighbouring mainland of Spain. 

Remains of Muscardinidse are known only from European 
deposits, and occur from Miocene times. Eliomys itself is repre- 
sented by E. poineli in the Lower Miocene of AUier, and by 
E. hainadryas in the Middle Miocene of La Grive St. Alban and 
Steinheim. Both species are of small size, and from the mandible 
do not show any close affinity to the remains which form the 
subject of this note, than which they would both appear to be 
already more highly specialised. In the British Museum there 
are two mandibular rami supposed to represent an Eliomys 
from the Pleistocene of Malta (B.M. 49342c and 49351) J. These 
are consideiably moie robust than in any recent member of the 
genus, and are found to resemble in shape the Mallorcan speci- 
mens and to be only slightly inferior in size, although the length 
of the alveolar area is considerably less. Unfortunately none of 
the cheek teeth ai'e preserved, and the formation of the roots can- 
not always be satisfactorily deduced from an examination of the 
alveolar cavities, though in one of these specimens (B.M. 49351) 
it seems that in the first and second molars the posterior roots 
had become confluent as in the genus now to be described. These 
specimens seem worth mentioning as suggesting a possible exten- 
sion of the known distribution of the Baleaiic dormouse. So far 
as I am aware, no other rodent remains have been described from 
the Balearics, although so long ago as 1855 De la Marmora 
wrote § that he had observed indications of an ossiferous breccia 
in the hill of Belver near Palma, in which he had seen a bone 
which seemed to belong to a Lagomys or to a rabbit. 

The following description of the most important specimens 
obtained may be taken as applying to the genus so far as known 
at present. It is proposed that the genus be known as 

Hypnomys, gen. nov. 

Skull, mandible, and limb-bones as in Eliomys but more robust ; 
interorbital region wide and anterior portions of frontals greatly 

* Ann. Mag. Nat. Hist. ser. 7, vol. xi. 1903, p. 494. 

t Proc. Zool. Soc. 1901, p. 41. 

X Cat. Foss. Manim. Brit. Mus. Pt. i. p. 225 (1885) (as Myoxus sp.). 

§ Mem. Accad. Sc. Torino (ser. 1), vol. xxxvii. 1855, p. 59. 



OF EXTINCT MUSCARDINE RODENT. 211 

expanded. The infraorbital foramen opens anteriorly and the 
outer wall of the infraorbital canal is very robust with a wide 
base. The anterior palatine foramina penetrate for some distance 
the palatal plate of the maxillas which forms the greater portion 
of the palate. The angle of the mandible is perforated. Dental 
formula i. J-, pm. ]-, m. |, molariform premolars and molar crowns 
sub-quadrate in shape with low transverse lidges. Upper molars 
with one large wide internal and two smaller external roots. In 
the first and second lower molars tlie two posterior roots may be 
confluent for the greater part of, or for their entire length ; the 
last molar has two anterior and one large posterior root. Tibia 
and fibula joined. 

Skull. — The skull is represented only by some few fragmentary 
specimens none of wliich show the posterior portion, that is to say 
behind the frontals. An isolated and imperfect auditory bulla 
(B.M. M 1 1 671) was obtained which agrees in form and in the shape 
and position of the meatus with the corresponding bone in Dyromys. 
So far as available specimens show, the skull in general resembles 
and may be compared with that of Eliomys, which for our present 
purpose will be taken in a broad sense and to include Dyromys. 
Even the smaller examples (those from Mallorca) are larger and 
comparatively, as well as actually, more robust than E. lusitanicus, 
which is the largest of the recent species of that genus. Viewed 
from above, the nasals are seen to be practically flat and about 
the same width throughout their length. The interorbital 
portion of the frontals is wide and smoothly flattened and is 
defined by sharp lateral edges ; a measurement taken at the 
narrowest point is nearly half the antero-posterior length of the 
nasals in Hypnomys, whereas in E. lusitanicits it is not quite a 
third. Anteriorly the frontals expand rapidly by a smooth 
swelling and attain to a great width — about twice that of the 
interorbital region — where they are joined by the nasals and 
the nasal processes of the maxilla?. This expansion is caused by 
the very large size of the olfactory cavities (PI. I. fig. 14). There 
is a fine but distinct ridge on the dorsal aspect of the maxilla at 
the upper root of the zygoma from where the skull narrows 
rapidly to form the snout. The premaxillse are i-obust, slightly 
inflated dorso-anteriorly as in Eliomys, and deep dorso-ventrally, 
the rostrum being stoxiter ' than in E. lusitanicus. The infra- 
orbital region is not very well preserved in any of the Balearic 
specimens, but sufficient is shown by two or three examples to 
demonstrate that in this respect Hypnomys shows no close 
resemblance whatever to either a typical Murine such as liattus 
or to typical Sciurines such as Sciurus or Xpa'us. Though showing- 
more general resemblance, this portion of the skull in the Balearic 
genus still diflfers somewhat from that of any other of the 
Muscardinidse with which I have been able to compare it ; in 
some respects it seems to agree with that of Leithia., though 
unfortunately all the anterior portions of skulls of this last in 



212 



MISS DOKOTHEA BATE ON A NEW GENUS 



the British Museum are either very imperfect or else considerably 
crushed and distorted. The infraorbital foramen opet)s anteriorly 
and, as well as can be seen in the type specimen of Hypnoiays 
mahonensis (PI. I. fig. 2), it has a greatest length of about 3 mm. 
and occupies .a median position between the two roots of the 
zygomatic process of the maxilla. The lachrymal foramen shows 
an extension of similar length, but ovoid in form, in front of the 
infraorbital foramen much as in Muscardimts, and without the 
lower lateral expansion seen in Eliomys. The outer wall of the 
infraorbital canal is very robust, broadly concave anterioi'ly, with 
a distinct outstanding point in front of its base, which is wide 
owing to the hinder border being produced outwards from above 
the first molar. 

The anterior palatine foramina start a short distance behind 
the incisors and penetrate the palatal plate of the maxillae for 
some distance, their extent being comparable with that obtaining 
in Eliomys. The palate (PI. I. fig. 1) is not completely preserved 
in any of the specimens, but can be seen to be wide and gently 
concave and chiefly composed of the palatal plate of the maxillfe, 
which extends posteiiorly about as far as the hinder border of 
the second molar. The palatal plate of the palatines is hardly 
shown but can certainly have formed only an insignificant 
portion of the palate. 

Text-fieure 1. 




Left mandibular rami of : — 

A. Leithia melitensis (B.M. 49342 D), 

B. Hypnomys morpheus (H.M. M 11097), 

C. Eliomys lusitanictis (recent specimen). X 1^. 

Mandible. — In general plan the mandible (text-fig. 1, B) is 
essentially as in Eliomys (text-fig. 1, C), from which it yet differs 
strikingly, although this diffei'ence is easier to observe than to 
describe. That of Eliomys may be s:iid to have an aspect of 



OF EXTINCT MLSCARDIXE RODENT. 213 

attenuation, its posterior portion particularly being very slight 
and the cheek-teeth closely crowded together. This applies also 
to the skull, though perhaps to a lesser degree. The mandible of 
the Balearic genus is altogether ixiore robust, and gives the im- 
pression of being possibly less highly specialised. The angle of 
the jaw is perforated, and also resembles in shape that of Eliomys, 
but compared with this last the coronoid process originates 
further forward and rises more abruptly ; it tapers to a slender 
point but is comparatively shorter and the space between it and 
the condyle less deeply excavated : this last also applies to the 
region between the condyle and the highest point of the angle, 
causing the hinder portion of the jaw to be more sohd owing to 
this comparatively greater extent of bone. The articulating 
surface of the condyle is strongly marked. The symphysial 
region and that between it and the cheek-teeth are likewise 
robust. The incisor extends considerably behind and rises above 
the cheek-teeth row, causing a marked protubex-ance on the out- 
side of the jaw at the base of the coronoid process, and the 
iirferior dental foramen occupies a correspondingly high and 
backward position. 

Teeth. — The dental formula is the same as that of other of the 
Muscardinid?e, that is to say, i. \, pm. \, m. |-. The incisors are 
of medium size, with the anterior band of enamel smooth and 
stained the charactei-istic orange-yellow colour. The upper 
incisor originates above and just in front of the premolar; its 
inner surface (PI. I. fig. 10) is flat and the outer gently rounded, 
this being also the case in the corresponding tooth of the lower 
jaw. Both are considerably compressed laterally with the antero- 
posterior thickness much greater than the lateral width, this being 
nearly double in the lower teeth. Their transverse sections (PI. I. 
figs. 11 k 12) are more or less elliptical in shape, thus differing 
from those for instance of Eliomys, Glis, and Leithia, in which the 
section forms practically an isosceles triangle with the anteiior 
face as the shorter base (PI. I. fig. 13). There is naturally a 
corresponding difference in the shape of the worn surfaces of the 
teeth. As Mr. Hinton has pointed out to me, these diffei-ences in 
the form and proportions of the incisors are of impoi'tance as 
indicating probable differences in the modes of life in these 
various genera. 

It is interesting to find that an examination of the microscopic 
structui-e of the enamel in the incisors of Hypnomys seems to 
bear out the conclusions independently arrived at from a general 
study of its remains. Mr. Thornton Cai'ter very kindly under- 
took the task of making this examina,tion, and allows me to 
include his report of his investigations, which is as follows : — 

" The structure of the enamel of the incisors in the specimens 
from Menorca and Mallorca is identical with that of Leiihia 
meliiensis. 

"The 'pattern' is distinctive and presents charactei^s which 
would seem to place it between Sciuridse and Myoxidee. In 

Proc. Zool. Soc— 1918, No. XVI. 16 



214 MISS DOROTHEA BATE ON A NEW GENUS 

longitiulinal sections the enamel rods (or fibres, as Sir John 
'J'omes designates them in his classical memoir) leave the surface 
of the dentine with a slight curve and then proceed outwards, 
slightly flexuous, at an angle of about 70° with the dentinal sur- 
face for about half the thickness of the enamel, where they bend 
sharply and abruptly to proceed in a straight course to the 
svu'face. 

" There is appearance of serrations of the margins such as is 
usually seen in the Dormice. 

" Thus the course of the rods is as in Myoxidse, whilst their 
form is as in Sciuridse, 

"In transveise section altei'iiate groups of rods cross one 
another at an angle of about 120°, and when half-way through 
turn sharply outwards and run paiullel out to the enamel surface 
Avhich they i-each at right angles, the pattern assumed resembling, 
though not identical with, that seen in a transvei'se section of the 
incisor of Eliomys." 

While the palate is almost uniform in width, the rows of cheek- 
teeth appear curved owing to their inner borders being practically 
in a straight line, so that all the differences in the sizes of the 
teeth, especially that of the premolar, alfect only the outer 
border of the rows, which thus have a curved outline. The plane 
of wear of the premolar is almost horizontal, while that of the 
molars lies at a considerable angle, the inner border being the 
higher ; this character will be refeired to again later. (Some 
scarcely worn upper molars are slightly concave and their enamel 
ridges incline towards being cuspidate, but in most of the specimens 
obtained the worn surfaces of the crowns are nearly flat. The 
molariform uppei- premolar is bluntly triangular in outline and 
the molars snbquadrate in shape, the second being very little 
larger and the third not much smaller than the first. In the 
above characters these teeth somewhat resemble those of Leithia, 
those of Eliomys differing markedly in the concavity and greater 
comparative width of their crowns. 

In the Balearic molars the ridges are low, and in those of aged 
individuals, of which a number were obtained, these become 
almost entirely woi^n away. In little-worn specimens (PL I. 
fig. 1) it can be seen that in each of the upper cheek-teeth the 
crown is crossed by four complete and one incomplete transverse 
ridges, with the addition of one or moi-e further incomplete 
ridges in the second and third molars. The second complete 
ridge in the first and second molars runs from the inner border 
in an anterior direction for more than half its distance before 
turning to reach the outer margin. The two posterior complete 
ridges run more or less parallel with the hinder border of the 
teeth, and it is in the space left between the complete ridges that 
the incomplete ones occur. The inner edge of the crown surface 
almost invariably remains unbroken, while the outer border is cut 
by the valleys between the ridges. In much worn teeth there 
is found a confluent valley alongside the inner enamel border of 



OF EXTINCT MU CARDINE RODENT. 215 

the tooth for the whole of its distance ; this condition of wear is 
also seen for instance in Leithia, Eliomys, and Xerus. Apart 
from size, the upper molars of Leithia chiefly differ from those of 
Hypnomys in their transverse ridges being considerably higher 
and more sharply defined, and in the complete ones in the first 
and second molars originating at the postero-internal border of 
the crown as already described and figured by Mr. Lydekker*. 

Of the lower cheek-teeth the premolar is the smallest and is 
obtusely triangular in shape ; the third molar is somewhat larger 
with a rounded posterior margin, although the crown has a 
tendency to become squarer in outline when much worn. The 
first and second molars are the largest of the series, almost equal 
in size and in shape quadrate, or with the antero- posterior length 
slightly greater than the width. The number of ridges appears 
to be much the same as in the upper cheek-teeth, though they 
become broken up earlier and into a greater number of incomplete 
ridges. In a moderately worn specimen (B.M. M 11674, PL I. 
fi,g. 6) the first and second molars seem to have three complete 
ridges, two distinct incomplete ones running half-way across the 
crown from opposite sides and a shorter one coming in from the 
antero-external corner, while the antero-internal area is broken 
up into several rather indistinct loops. The third molar shows 
four complete ridges and two small accessory ones. In the case of 
aged individuals the confluent worn surface is of course along the 
external border of the crown, it being on the internal side in the 
upper cheek-teeth. In less worn specimens this external enamel 
border is interrupted at two points, and the enamel between and 
on either side of these channels, owing to its being slightly folded 
over, is at this stage of wear more massive than in other parts of 
the ridges. 

The number, position, and conformation of the roots of the 
cheek-teeth in rodents form a subject of considerable interest and 
importance, and it is one that has received a good deal of atten- 
tion from investigators, notably Schlosser t and, later. Dr. Foi-syth 
Major J. The specimens under discussion are no exception in 
this respect, for their molar roots are unlike those of any Pleisto- 
cene or recent genus with v/hich I have been able to compare 
them with the single exception of Leithia, and that only in the 
case of the upper molars. It may be mentioned that I have not 
found this character previously noted in the descriptions of the 
Maltese genus. The alveolar formula can be most easily realised 
by reference to the accompanying text-figure (text-fig. 2), but at 
the same time it should be borne in mind that such a formula 
alone is of very little value for purpoess of comparison. The 
pi^esent examples are a case in point, as, for instance, the first and 
second upper molai's of Eliomys and Glis are three-rooted, as in 

* Pi-oc. Zool. Soc. 1895, pp. 861-3, text-fig. 1. 
t ' Pakeontographica,' Band xxxi. 1885. 

X See, for instance, the following : " On Fossil Dormice," Geol. Mag. Nov. 1899, 
]>. 493, and " Ou some Miocene Squirrels," Proc. Zool. Soc. 1893, pp. 179-214. 

IG* 



216 MISS UOROTUKA BATE ON A NEW GENUS 

Hyjmomys and LeAthia, but an examination of tlie roots them- 
selves show them to be essentially different. 

In Hypnomys the upper premolar has three long and rather 
stout roots, which terminate in blunt rounded ends(B.M. M 11658, 
PI. I. fig. 3). The anterior one takes a forward direction and is 
the shortest of the three, the other two project in a parallel 
direction from the hinder border of the tooth and may be con- 
fluent for a varying distance, but apparently not for more than 
half their length. This description is based on examples of 
Hypnomys mahonensis, but the alveolus in a Mallorcan specimen 
indicates a similar formation. Unfortunately there has been no 
opportunity of examining any milk-teeth in the case of either 
this genus or Leithia. 

Text-figure 2. 

A. B. 

*>g f nm . nm "^ e nm ® rim 



^ II/V3 



@9 



>a 



(/. alveoli, L.olveoli. U. alveoli. L. alveoli 

Diagram of alveoli of cheek-teeth of A. Sypnomi/s and B. LeitMa. 

The upper molars each have three roots, one large internal and 
two small external ones, a number commonly found in many 
rodents. The distinctive feature of the Balearic teeth lies in the 
construction of the large internal roots : in Eliomys, Glis, and 
also in other forms such as Xenos and Sciums, which have the 
same alveolar formula, these are single, rather flattened conical 
roots projecting from the centre of the border of the crown which 
they support. In Hypnomys the internal root is quite different, 
this being clearly portrayed in the accompanying illustrations. 
Fig. 3 (PI. I.) shows the latero-internal view of the first and 
second molars of H. mahonensis : it will be seen that there is a 
slight consti'iction at the base of the crown where the enamel 
ends, and from there the confluent root continues practically of 
the same size for its whole length, which is a little less than twice 
the antero-postei-ior length of the crown-surface. In the figured 
specimen the first molar measures 4 mm. from the crown-surface 
to the tip of the root, while the antero-posterior length of the 
crown is veiy little more than 2 mm. In the second molar the 
loot is very slightly shorter and not rounded at its apex, but this 
n;ay perhaps be partly due to damage in developing the specimen. 
The internal root of the third molar is exposed in the type 



OF EXTINCT MUSCARDINE RODENT. 217 

specimen of H. mahonensis (B.M. M 11657), in which it is seen 
to be similar to that of the other two. In some specimens there 
is a slight groove along the centre, indicating the compound 
nature of the roots ; the ends are closed in all examples examined. 
The slight intero-external width is well shown in the section 
(B.M. M 11661, PI. I. fig. 4) and in the posterior view of the 
second upper molar (B.M. M 11660, PI. I. fig. 5), in which it can 
also be seen that there is a cei'ta,in amount of curvature in the 
direction of the compound roots. They are considerably longer 
than the external roots, although, owing to the slope of the 
crowns, they do not penetrate much deeper into the alveolar 
ca\^ity. 

As already mentioned, the upper molar roots of Leithia are 
similar to those of Hypnomys^ the only other genus, among many 
examined, showing a closely similar condition of root structure is 
Trechomys, In the British Museum there are two fragments 
of the upper jaw of T. platyceps (B.M. M 1627) from the Phos- 
phorites of Caylux, containing the premolar and first molar, the 
I'oots of the latter being as in Ilypnomys except that the broad 
inner root is perhaps not quite so wide compared with the crown. 
It must be mentioned that in Trechomys the roots of the premolar 
are the same as in the molars, whicii is of coui^se not the case 
in either Leithia or the Balearic genus. The molar roots of 
Theridoriiys are somewhat similar but are accompanied by much 
more hypsodont crowns. 

The roots of the lower cheek-teeth of Hypnomys (B.M. M 11673, 
PI. I. fig. 7) show a very considerable amount of variation botli 
as regards their length and conformation. The premolar may 
perliaps be said to have two roots, but these ai-e confluent for 
the greater part of their considerable length, diverging at a vary- 
ing but never very great distance from their apices, the anteiior 
of which is slightly the longer of the two. The upper, posterior 
portion of the root is wide and flattened, being very evidently the 
result of the fusion of two roots. In some specimens (as in B.M. 
M 11678) there are clearly three confluent roots. 

In the first and second molars the two anterior roots are 
usually long, equal in size, and separate though not very widely 
divergent! In the posterioi' roots the length and antero-posterior 
thickness are about the same as in the anterior pair, so that 
viewed laterally little or no difference is observable (PI. I. fig. 7). 
But viewed from behind they are seen to be confluent transversely 
for either the whole or three-fourths of their length (PI. I. 
tigs. 8, 9, B.M. M 11675-6); in the latter case the stout terminal 
portions of the roots may diverge considei'ably. The resembla,iice 
to Hypnomys seen in the roots of the upper molars of Leithia 
does not hold good for those of the lower jaw, for in the 
Maltese genus the first and second molars each have four roots, 
entirely separate from each other for their entire length. In 
Trechomys, however, the lower molar roots seem to a.gree with 
those of the Balearic genus with the pos&ible exception of the 



218 MISS DOROTUEA BATE ON A NEW GENUS 

tliiid moliir. Schlosser gives the nnml)er of roots for tliis tooth 
as three, hut judging f loiu the alveohis of a specimen of T. platyce])s 
in the British Museum (M1627) it appears that there may some- 
times be two large, transverse roots only. 

The last lower molar in Hypnomys has three roots, two anterior 
slender ones and a large, backwardly projecting posterior one. 
This condition is very similar to that obtaining in the correspond- 
ing tootli of Leithia and the recent Eliomys, though in the latter 
there is a more marked constriction immediately below the crown. 

Limb-boiies. — Very little of the skeleton was obtained, but a 
few femorn, a tibia, and fragmentary portions of humeri and 
other bones, chiefly from a Mallorcan cave-deposit, show characters 
which also indicate that the genus should be included in tlie 
Mnscardinidje. The ulna is very similar to that of Eliomys, as is 
also the femur, except that the third trochanter seems to be 
slightly less prominent and less sharply separated from the great 
trochanter. As contrasted with the Hystricidpe, SciuridtB, and 
Anomalui'idfe, which have the tibia and fibula free, a charac- 
teristic of the Muridse and Muscardinidfe is the joining of these 
two bones for a third or moi'e of their distal portions. It may be 
woi'th noting that in Rattus {Epiinys), for instance, the fibula 
at its distal extremity is once more separated though closely 
adpressed to the tibia. On the other hand, in Eliomys q-icercimis 
the tibia and fibula continue completely ankylosed at their distal 
ends, and it is with this latter that the Balearic specimens agree, 
as Leithia likewise does in this respect. These bones are joined 
for more than a third of their total length in the thi'ee last- 
mentioned genera. In Hypnomys the tibia is a stiaighter bone 
with a less well-developed cnemial crest than in Eliomys. 

The above are the chief characters of all the Balearic specimens, 
and it will only be necessary to add a brief note and a few 
measurements in the two species from the difterent islands. 

Tiie species from Menorca may be named 

Hypnomys mahonensis, sp. n., 

aiid regarded as the type species of the genus, distinguislied from 
the Mallorcan species by its greater size and diiterent habitat. 
Fm-ther diftereuces would probably be apparent were a larger 
amount of material available. All the specimens 'Were obtained 
from fissures in the Miocene limestone, and were in one or two 
instances associated with i-emains of Myotragus and Tesiudo 
gyninesicus. Some imperfect lacertilian jaws also occurred, and 
tliese have been very kindly examined by Mr. G. A. Boulenger, 
F.R.S. : they prove to be those of a Lacerta of the viridis-ocellata 
gi-oiip 'and a species of Chcdcides. Although representatives of 
both these occur commonly in the Mediterranean region, neither 
are found in Mallorca or Menorca at the present day. 

Owing to the fragmentary state of the specimens not many 



OF EXTINCT MUSCARDINE RODENT. 219 

measurements can be given, but the following are a few. The 
length of the nasals is about 18 mm. in the single example in 
which these bones are completely preserved, and the interorbital 
width of the frontals is 8 mm. The base of the outer wall 
of the infraorbital canal measures 6 mm., and the incisive 
foramina are approximately 8 mm. in length. In the type 
specimen (PI. I. fig. 1) the complete row of left upper cheek-teeth 
is almost 10 mm. in length, this measurement being 14 mm. in a 
corresponding example of Leithia, and barely 7 mm, in one of 
Eliomys lusitanicus. In II. mahonensis the width of the palate 
between the first molars is not quite 6 mm., in Leithia it is little 
more, being 6'5 mm. 

The length of the lower cheek-teeth series is about 10 mm. ; 
as already mentioned the length of their roots is very variable : 
in one specimen having teeth with moderately worn crowns 
(B.M. M 11673, PI. I. fig. 7) the first molar has a crown-width 
of 2-5 mm. with a root-length (measured from the crown-surface 
to root-tip) of almost 5 mm. In the third molar in the same 
specimen the greatest length of root is 3 mm., the large posterior 
root attaining an antero -posterior width of 1"5 mm., while the 
antero-posterior length of the crown is 2'5 mm. In an example 
of a left lower second molar (B.M. M 11676, PL I. fig. 9) with a 
less worn crown than the above the crown-width is 2 mm., and 
the greatest root-length 3 mm. 

It is proposed that the species from Mallorca be known as 

HyPNOMYS MORPHEUS, Sp. n., 

characterised by its smaller size and difierent habitat from 
H. mahonensis. The few specimens of jaws and limb-bones by 
which it is represented were obtained from cave-deposits in 
Mallorca, and were found associated with remains of Myotragus, 
and in one case with a few mandibular rami and limb-bones of 
Apodemtts, which still occurs plentifully in the island. In size it 
appears to agree with the larger forms of Glis. The type speci- 
mens (B.M. M 11695), consisting of the anterior portion of the 
skull with the incisors and right molars and two mandibular 
rami, are believed to have been associated and fell apart on being 
sepai-ated from the earthy matrix in which they were preserved. 
The base of the outer wall of the infraorbital canal measures 
5 mm., and the antero-posterior length of the molar row is 6 mm. 
The length of the lower cheek-teeth series is 8 mm. In the 
upper incisor (B.M. M 11696) shown in fig'. 10 (PI. I.) the antero- 
posterior width is 2-5 mm. while the thickness is 1*5 mm. ; in a 
specimen of the lower incisor the antero-posterior width is 2 mm., 
with a thickness of barely more than 1 mm. 

The greatest length of a right tibia, wanting its proximal 
epiphysis, is 41 mm., the distal 18 mm. of which are joined to the 
fibula. A femur has a total length of 55 mm., that of a right 
ulna being 36 mm. 



220 MISS DOROTHEA BATE ON A NEW GENUS 



Summary and Conclusions. 

The above description may be very briefly sumiTiarised as follows. 
IJypnomjjs is represented by two species known by i-eninins from 
the two largest islands of tlie Balearic gronp and is consideied 
to belong to the Muscardinidpe, though seeming to be a very 
distinct form not closely related to any other at present known. 
Its dental and osteological characters so far observed not only 
seem to show that Ilypnomys belongs to the Mnscardinidse but 
also appear to afibi-d further support to the opinion, now very 
generally held, that this group cannot be included in either the 
so-called Sciuromorpha or Myomorpha and lending additional 
weight to the argument against employing these terms. This 
seems to hold good also for the modern representatives which 
have been briefly described by Mr. Miller * as "... mostly 
arboi'eal animals with habits and aspect somewhat intermediate 
between mice and squirrels. . . ." 

The Balearic genus may be compared with the recent Eliomys 
and the extinct Leithia from the Pleistocene of Malta. It agi-ees 
with the former in genei-al plan of skull, lower jaw, and limb- 
bones, and it may be noted that there is also a perforation in the 
angle of the lower jaw. It differs in a number of points, among 
which the following may be cited : — In the pattern of the molar 
crowns, which are very slightly concave and lack the marginal 
cusps, both distinctive characteristics of Eliomys. In Hyj^nomys 
the worn surfaces of the molars are subquadrate in shape, whereas 
in the recent genus the width is distinctly greater than the 
antero-posterior length ; the alveolar formula differs in the two 
genera, and the large inner root in the upper molars of Hypnomys 
is quite distinctive. This root- structure was evidently attained 
by a process of simply filling up the space between two parallel 
roots, whereas the single conical root seen in the Scduridse and 
others would seem to be arrived at by the drawing together of 
the apices of the two roots with a similar filling up of the 
intervening space. It would be interesting could one arrive at 
a reasonable hypothesis to account for this root-formation in 
Hypnomys and Leithia, for presumably it must have been acquired 
to meet a special stress or strain consequent on some peculiaiity 
of diet or mode of life. 

Ilypnomys agrees with Leithia in its squared molars and in 
the large, confluent and single inner roots of the upper molars, 
but differences are seen in the former having a perforation in the 
angle of the jaw, which is also of a different shape (see text-fig. 1), 
in the crown pattern of the molars, much lower enamel ridges, 
and in the roots of the lower molars. 

It has also been noted that in the roots of both upper and 
lower molars, but not premolars, Uypnomys shows a striking 
resemblance to Trechomys. However, it is not for a moment 

* ' Cat. Mammals of West. Envope,' London 1912, p. 549. 



OF EXTINCT MUSCARDINE RODENT. 221 

suggested that this similarity, which is apparently unaccompanied 
by other points of resemblance, indicates any close affinity. On 
the contrary, it would seem that these three genera each exhibit 
in this character what would have been a stage, and that a very 
early one, in the history of the evolution from the low-crowned 
four-rooted towards the hypsodont and rootless or semi-rootless 
molar. The cheek-teeth of Theridomys also seem to suggest the 
possibility of this result being attained without the necessity of 
including a stage having a large conical inner root in the upper 
molars accompanied by a somewhat triangular-shaped crown, as 
seen in some of the Sciuridse {Xerus for instance) and which 
Dr. Forsyth Major has fully described in his invaluable paper on 
the Miocene Squirrels*. 

HypnoviTjs is an interesting addition to the extinct fauna 
of the Balearics so far known by Myotragus and Testudo 
gymnesicus, both forms totally distinct from the present-day 
fauna of the islands. Owing to the small amount of material 
available, the question as to the precise age of these Balearic 
dfeposits and their contents seems a difficult, if not impossible, 
one to answer at present. Lately this matter has been the 
subject of some interesting remarks by Prof. M. Boulet, who 
seems inclined to consider the deposits of earlier age than 
Pleistocene. Seeming to support this view are the absence of 
human remains and the character of the chief remains found, i. e. 
Myotragus, Hypnomys, and a giant Testudo. The occurrence of 
remains of this last in the limestone fissures of Menorca certainly 
suggests that there must have been great changes of climate and 
vegetation, and that there have been considerable alterations in 
the land surface is indicated by extensive stretches of sheets of 
stalagmite now exposed on the weathered surface of the Miocene 
limestone in both Mallorca and Menorca. 

On the other hand, it should be realised that not much weight 
should be placed on the absence of human remains owing to the 
fact that not a single one of the ossiferous deposits was found to 
be in an undisturbed condition, all the upper layers in which such 
remains might be expected to occur having been entirely destroyed. 
Another point to be borne in mind is that in the Mediterranean 
region several species now extinct have been proved to have per- 
sisted until the time of man's occupancy of these particular areas. 
The well-known Prolagits, remains of which were found in 
Neolithic settlements in Corsica by Dr. Forsyth Major, is a case 
in point. Another somewhat similar example is that of a small 
deer, AnaglocMs, whose remains occur abundantly in some of the 
cave-deposits of Crete, while a number of its antlers were found 
by Sir Arthur Evans in a shrine in the famous Minoan Palace 
of Knossos. 

I should like to take this opportunity of expressing my grateful 
thanks to the Trustees of the Percy Sladen Memorial Fund for 

* Pi-oc. Zool. Soc. 1893, pp. 179-214. 

t ' L'Autlii-opologie,' tome xxviii. (1917) pp. 160-3. 



222 A NEW GENUS OF EXTINCT MUSCAUDINE RODENT. 

enabling me to visit Menorca. Also to Dr. A. Smith Woodward, 
F.K.S., for continued kindness in giving me every facility for 
working in the Geological Department of the British Musei;m 
(Nat. Hist.), to Mr. M. A. C. Hinton for kind advice and for 
looking over some of my material, and to Mr. J. Thornton Carter 
for his vahiable investigations into the enamel structure in the 
teeth of Hypnomys and other genera. 



EXPLANATION OF PLATE L 

Fig. 1. JTypnomys wahonensis. Tj'pe-specinien (P.M. M 11657). Palate showing 

tlie left premolar and molars and right M'. X 4. 
Side view of same specimen showing anteovbital region. X 3. 
Lateral view of right upper premolar and 1st and 2nd molars, 

showing the large confluent roots of the latter (B.M. M 11658). 

X3. 
Transverse section of left M^ (P.M. M 11661). X 4. 
Posterior view of right M2 (B M. M 11660). X 4. 
Crown view of right lower molars (P.M. M 11674). X 4. 
Lateral inner view of right lower cheek-teeth (P.M. M 11673) . 

.X3. 
Right Ms (P.M. M 11675) showing posterior roots. X 4. 
Left Mo (P.M. M 11676) showing posterior roots. X 4. 

10. a. morpheiLS. Right upper incisor (P.M. M 11696), inner view. X 3. 

11. a. malwnensis. Transverse section of upper incisor (P.M. M 11662). X 4. 

12. Do. Transverse section of lower incisor (P.M. M 11677). X 4. 

13. Leithia melUensis. Transverse section of upper incisor (P.M. M 49345 a). 
X4. 

14. if. malionensis. Dorsal view of interorbital region of skull (P.M. M 11659). 
X2. 



2. 


Do. 


3. 


Do. 


4. 


Do. 


6. 


Do. 


6. 


Do. 


-7. 


Do. 


8. 


Do. 


9. 


Do. 



ON HERMAPHRODITISM IN A LIZARD. 223 

12. A Case of Hermaphroditism in a Lizard, Lacerta viridis. 
By Noel Taylek, B.Sc. (Lond.).* (From the Zoo- 
logicnl Departmentj University of London, University 
CoHege.) 

[Received April 23, 1918 : Read May 28, 1918.] 
(Text-figures 1-3.) 

The specimen, the nrogenital system of which is described in 
the following pages, was placed in my hands by Prof. J. P. Hill, 
P.K.S. It turned up in the course of class-work in the Senior 
Laboratory in the College, and its abnormal condition being 
oliserved, it was fixed in corrosive sublimate and preserved for 
detailed examination. The specimen presents certain features 
of interest which, it is believed, are worthy of being placed 
on record. 

I wish to express my thanks to Prof. Hill for his advice and 
assistance in the preparation of these pages. 

I. Descriptive. 
a. General Morphology. 

When this specimen came into my hands, it was in a partly 
dissec^ed condition, the greater portion of the abdominal viscera 
having been removed. 

The Fat Bodies were present and well developed, the right 
being rather larger than the left. They are not represented, 
however, in text-fig. 1, which gives a general view of the 
urogenital system, since in sihi they obscured the more anterior 
portions of the oviducts. 

The testes (text-lig. 1, E.T. and L.7\) were well developed and 
suspended in the folds of the longitudinal mesorchia (Afes.). 
The right was placed somewhat more anteriorly and was rather 
larger than the left, the dimensions of the former being about 
•9 cm. X '4 cm., and of the latter "65 cm. x '3 cm. 

Both gonads seem normal in shape apart from the remarkable 
stalked outgrowths on each (Ov.). These on section were found 
to contain ova, and the gonads may therefore properly be 
designated ovotestes. The right gonad, it will be seen, possessed 
two of these spherical ovarian appendages, each joined to the 
dorso-lateral border of the testicular portion of the organ by a 
well-marked stalk; the more anterior was further sub- divided 
into two by a median constriction. 

The left gonad also possessed two outgrowths, the surface of 
the larger being subdivided into five or six hemispherical pro- 
jections. The epididymes were well developed (L.Up., R.Ep.) 

* Coinmunicated by Prof. J. P. Hill, D.Sc, P.R.S., F.Z.S. ' 



224 



MR. NOEL TAYLER OX 

Text-fii>nie 1 






rt: 




JLaeerta virirUs: dissection of a hermaplirodite animal. 
IS.vplanation of the lettering. 

Mes. Mesorcliium. 



B.Lq. 


Bvoad Ligament. 


Bl. ' 


l^Lidder. 


d.aorf. 


Doreal Aorta. 


L.Cop. 


Left Coitulatory Organ. 


L.Ep. 


Lett Kpididviiiis. 


B.FI. 


Leit F.inuei. 


L.M.D. 


Lett Oviduct. 


JLnq. 


Luii^'. 


L.T. 


Lett Testis. 


L.v. 


Liver. 



Ov. 


Ovary. 


B.Coj>. 


Riiilit Copnlatory Organ 


B.Ep. 


Right P]piilidymis. 


B.K. 


Right KidnP3% 


li. L. 


Muscular Band. 


R.M.D. 


Riglit Oviduct. 


B.T. 


J{iglit Testis. 


Bt. 


Rectum. 


Sp. 


Spleen. 



HERMAPHRODITISM IN A LIZARD. 225 

and situated in the broad ligament parallel to but separate from 
the testes. They were not attached, as is normally the case, to 
the inner border of the testes, and vasa efFerentia were conse- 
quently absent. Posterioi-ly, the epididymes passed into the 
vasa deferentia, and these, having united with the ureters, opened 
into the posterior division of the cloaca by the urogenital papillas. 

The two kidneys were apparently normal, each consisting of 
an anterior and posterior lobe (in the figure, only the right 
kidney, U.K., is indicated by reference letters). 

The copulatory organs (li. Co^J., L. Cop.) were present as in 
the normal male lizard. 

The oviducts {R.M.D., L.M.D.) were developed for about 
a third of their lengths. Each opened into the body-cavity by a 
well-developed funnel (only the left, L.FL, is indicated by 
reference letters in the figure), and behind that was continued 
into the duct, the plaited glandular walls of which are seen in 
the outer border of the broad ligament (B.Lg.). The right 
oviduct attained the greater complexity as in the normal female. 

Passing down from the posterior tip of the ccelomic funnel on 
each side, on the extreme outer border of the broad ligament 
was a narrow but well defined ribbon-like muscular band which 
continued right back on each side to the cloaca.. Similar bands, 
which were at first taken for the oviducts, were referred to by 
Howes (5) as the round ligaments, and he thus describes them 
in his specimen : " From this (the oviducal aperture) there passed 
back a ribbon-shaped muscular band which skirted the free edge 
of the broad ligament, remindful of the round ligament of 
mammalian anatomy. This structure was wholly absent on the 
side destitute of an oviducal vestige, as indeed it is in the normal 
male. Its development is correlative with that of the oviduct." 

b. Histology. 

Transverse sections were made of the gonads and stained with 
hasmatoxylin and eosin. The main body of each gonad was found 
to consist of normal testicular tissue, i. e. seminiferovis tubules 
with an interstitial sti^oma, the lining cells being in active 
mitotic division (text-fig. 2, s.t.). 

The stalked outgrowths were found to consist of ovarian tissue, 
the bulb-like extremities containing large and fully-grown ova. 
A section of the gonad through the nuclear plane of the most 
anterior ovum is somewhat diagrammatically represented in 
text-fig. 2. 

The large yolk-laden ovum {ov.) is, it will be seen, surrounded 
by the relatively thin follicular epithelium {foil, ep.), outside 
this is a fibrous layer continuous with that sheathing the 
testicular portion of the gonad. In the stalk region all the 
normal histological elements of the ovary are represented. 
Outside is a layer of cubical epithelial cells continuous with 
the peritoneal epithelium, while the main mass of the stalk is 
formed of a loose stroma of connective tissue, contained in which 



•226 



MR. NOEL TAYLER ON 



are numbers of young follicles of various ages, tlie youngest 
being nearest to the testis. The connective-tissue body of the 
stalk is confluent with the tunica albuginea of the testicular 
portion of the gonad, which is somewhat thickened in the i-egion 
of junction. 

The sections of the oviducts revealed in their anterior portions 
a typical structure. They are lined with a well developed 
ciliated and glandular epithelium, the lumen of the duct being 
filled witli the coagulated secretion of the gland-cells. Sperma- 
tozoa were not detected. 

Text-figure 2. 




fo/l. ep. 




Lacerta viridis, hermaphrodite animal: sections through gonad. 

ov.; ovum. 

foil. ep. ; follicular epithelium. 
s. t. ; testicular stroma. 



The appearance of the epididymis was quite normal, but no 
spermatozoa were found in the lumen of the vas deferens, and 
examination of the sections failed to reveal the pi-esence of 
vasa efierentia through which the spermatozoa norxnally pass 
from gonad to epididymis. 



HERMAPHRODITISM IN A LIZARD. 



227 



111 the dissected specimen two or three fine tubule-like filaments 
were seen to pass from the posterior ends of the oviducts to 
the anterior extremities of the corresponding epididymes ; these 
were best marked on the left-hand side (see text-fig. 1). These 
filaments were carefully examined in the sections. The epi- 
thelial lining of the oviduct was found to end posteriorly in 
a cul-de--sac, as did that of the epididymis anteriorly. The 
"filaments" in section appeared as actual tubular passages 
devoid of any epithelial lining and running in the substance of 
the broad ligament. They commenced at the extreme anterior 
tip of the epididymis, and while two of them ended blindly, the 
third ran up as far as the posterior extremity of the oviduct, its 
lumen being continuous with the spaces in the mesentery in 
which were contained the glandular portion of the oviduct. 



Text-figure 3. 







Lacerla viridis, herraaplirodite animal ; 
kid. ; kidney tissue. 



thec.folL 

section through kidney. 



foil. ep. ; follicular epithelium. 
theo.foU. ; theca folliculi. 

The morphological interpretation of these spaces would seem 
a problem of some dilficulty. Being devoid of epithelial lining 
and making no connection with the lumen of the epididymis, one 
would hardly seem justified in regarding them as rete tubules 
proper, while the close relationship of one of them with the 
oviduct also seems anomalous, on this view. 

The condition which Howes (5) describes in one of his 
specimens seems of some interest in this connection. " In one 
instance," he says, " I discovered an interesting modification of 
the condition recorded by Leydig. There was buried up in the 



228 MR. NOEL TAYLER ON 

peritoneum in a line with the head of the epididj'iuis a dehoate 
filainent which, while it answered in every respect to the 
rudiment described by him, instead of ending abruptly and 
blindly became suddenly en]a,rged, opening into the body-cavity 
by a wide-mouthed funnel-shaped extremity, identical with that 
of the oviduct, and lined by a ciliated epithelium." 

This without doubt seems the description of a vestigial oviduct, 
yet, posteriorly, according to his figiu^e, it appears to arise from 
the antei'ior tip of the epididymis. Unfortunately he does not 
seem to have investigated the precise nature of its relations to 
this organ. 

The sections through the kidneys reveal the presence on the 
dorsal portion of one of them of an embedded mass of almost fully 
grown ova. Text-fig. 3 is a semi-diagrammatic representation 
of one of the sections in which five ova occur. It will be seen 
that the mass of ova lies actually embedded within the kidney 
tissue (kid.). Each ovum is surrounded by a layer of follicular 
cells (Jhll. ep.), while externally to this and separating it from 
the kidney substance is a thin fibrous layer (thee, foil.) presum- 
ably representing the theca foUiculi. 

II. Discussion. 

'No instance of complete hermajdiroditism in the Lacertilia 
seems yet to have been put upon I'ecoi'd, though cases of the 
more or less complete developmeut of the Miillerian Ducts in 
adult male lizards have been described. 

Leydig in 1872 (7) described the persistence in the males of 
Lacerta agilis of the Miillerian Ducts as small blind and con- 
voluted tubules, while Braun in 1877 (2) noted the development 
of rudimentary Miillerian Ducts in the 3'oung male of the same 
species, making no mention, however, of its presence in the 
adult form. 

In 1887 Howes (5) published a brief but important paper, 
" On the vestigial structui'es of the reproductive apparatus in 
the male of the Green Lizard " (Lacerta viralis). One of his 
specimens was a male lizard in which both the oviducts were all 
but full}^ developed, while in another the oviduct was fully 
developed on one side. 

In thirteen out of twenty-five specimens examined certain 
segments of the oviduct were well developed, the other portions 
being onl}^ represented as delicate filaments, thus giving a series 
of conditions analogous to those described by Matthews for the 
male toad. 

In 1893, Hill (4) published an account of the persistence of 
vestigial Miillerian Ducts in the full-grown male of an Australian 
lizard, Amj^hibolurus muricatus ; while two years later, in 1895, 
Jaquet (6) described the presence of Miillerian Ducts identical 
with those of the normal female in an adult individual of 
Lacerta ayilis. 



HERMAPUUODITISM IN A LIZARD. 229 

All the above mentioned cases are, it will be noted, pie- 
doininantly male, indeed as far as their gonads are concerned 
wholly male, for in no case is any reference made to the 
presence of ova or ovarian tissue. 

In this respect the specimen described in the present paper 
stands in striking contrast to previously described cases ; more- 
over, while it is distinguished by the presence of well-marked 
ovotestes, it must have been physiologically sterile. 

Among lower forms the occurrence of well-developed Miillerian 
Ducts seems often to be accompanied by the existence of an 
ovotestis. Cases of this kind have been described by Marshall 
in the Frog (8), but Fantham (3) seems to have been the first to 
record a case of true hermaphroditism in the Reptilia. The 
specimen of Testudo grceca described by him possessed well- 
developed oviducts, the lumen of each being continuous through- 
out. Of the two gonads the right was a typical testis, on the 
ventral surface of the left however was a "conspicuous yellow 
egg." On section another was found developing in its proximity, 
while " a few groups of bodies resembling developing ' ovarian 
ova' were seen scattered in separate groups (follicles) among 
otherwise testicular tissue, more especially near the periphery of 
the anterior portion of the gonad." Epididymes, vasa efferentia 
and vasa deferentia were present as in normal specimens, the 
former being rather large. 

It seems a point worthy of note that the development of the 
oviducts in the cases referred to above, viz., those of Howes, 
Hill, and Jaquet, is much more complete than in the subject of 
tlie present paper; in all these three cases the oviducts were 
developed throughout their whole length and opened into the 
cloaca, yet in none of these cases is any mention made of the 
presence of ova or ovarian tissue. In the present specimen, on 
the contrary, numerous ova occur, though only the anterior 
thirds of the two oviducts are fully developed. 



Literature referred to. 

1. ]^ouRNE, A. G. — " On certiuu abnormalities in the Common 
Frog; i. The occurrence of an ovotestis." Q. J. M. S. xxiv. 

3. BiiAUN. — " Diis urinogenital Syst. der einheimischen Ilep- 
tilien." Arbeit, aus dem zool.-zootomisch. Instit. WUrzburg, 
vol. iv., 1877-8. 

3. Fantham., H. B. — "On Hermaphroditism and Vestigial 

Structures in the Reproductive Organs of Testudo grceca." 
Ann. Mag. Nat. Hist. xvi. 1905, p. 120. 

4. Hill, J. P. — "Note on the presence of Vestigial Miillerian 

Ducts in a full-grown male lizard {Amphiboluriis muricatus) ." 
Proc. Linn. Soc. N.S.W. vol. viii. (Series 2ndX Sept. 27th, 
1893. 
Proc. Zool. Soc— 1918, No. XVII. 17 



230 ON HERMAPHRODITISM IN A LIZARD. 

5. Howes, G. B, — " On the Vestigial Structures of the Repro- 

ductive Apparatus in the Male of the Green Lizard." Jouru. 
Anat. & Phys. xxi. pp. 185-9, 1887. 

6. Jaquet, M., (1895). — "Note sur un cas d'hermaphroditisme 

incomplet observe chez le Lacerta ayilis." Bibliogr. Anat. 
(Paris-Nancy) iii. p. 267. 

7. Leydig. — "Die in Deutschland lebenden Arten der Saurier." 

Tiibingen, 1872. 

8. Marshall, A. Milnes. — " On certain abnormalities of the 

Reproductive Apparatus of the Frog." Journ. Anat. & 
Phys. xviii. pp. 121-144. 




P Z.S. 1919. WOODWARD, Pl.I. 



2x6 



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G.M.Woodward deLetliti. 

1. CROSSORHINUS JURASSICUS. 

2, 3. PROTOSPINAX ANNECTANS. 



Hufcii imp. 



ON TWO NEW ELASMOBRANCH FISHES. 231 



13. On two new Elasmobrancli Fishes (Crossorhinus juras- . 
sicas, sp. nov., and Protospinax annectans, gen. et sp. 
nov.) from the Upper Jurassic Lithographic Stone o£ 
Bavaria. Bf Arthur Smith Woodward, LL.D., 
F.R.S., V.P.Z.S. 

[Received May 23, 1918 : Read June 11, 1918.] 

(Plate I.) 

Most of the modern groups of Elasmobranch fishes seem to 
have arisen during the Cretaceous period, but some are of still 
older date, and a few interesting types are represented by well- ' 
preserved fossils in the Upper Jurassic lithographic stone of 
Bavaria, Wiirtemberg, and France. Two remarkable new 
examples of these early forerunners of the existing fainia have 
lately been identified in the British Museum, one apparently 
indistinguishable from an existing genus, CrossorJdvns {or Orecto- 
lohus), the other evidently of a new genus and family closely 
related to the Spinacidse. 

Family CROSSORHINID^. 

Crossorhinus jurassious, sp. n. (PI. I. fig. 1.) 

specific Characters. — Head gently rounded in front ; length of 
head and trunk about equal to that of the tail. Three pairs of 
flinging dermal lappets, all undivided, the first extending along 
the sides of the front half of the head, the next pair diminutive, 
and the third pair largest, extending along the sides of the 
branchial region. Pectoral fins rounded, relatively large, ex- 
tending nearly as far back as the origin of the pelvic fins, which 
are also rounded and about two-thirds as wide as the pectorals. 
Dorsal fins rather small and apparently nearly equal in size ; the 
first dorsal arising opposite the hinder limit of the pelvic fins* the 
second ending in advance of tlie much smaller anal fin, which is 
close to the lower lobe of the caudal. Body and fins covered with 
very fine shagreen, of which some granules between the pectoral 
fins have a fiuted sculpture. 

Description of Type Specimen. — The fossil, which is shown of 
the natural size in PI. I. fig. 1, is exposed in its anterior half 
from below, in its caudal half from the side. The snout is short 
and bluntly rounded, and the rami of the jaws are vaguely seen, 
meeting in an acute angle at the symphysis, where there are 
remains of a cluster of very slender, smooth, pointed teeth. The 
branchial region is relatively long, but the branchial arches are even 
more obscured by the crushed shagreen than the jaws. A single 

17* 



232 DR. A. SMITH WOODWARD ON 

pair of large dermal lappets (III), widest in front and graduMlly 
narrowing backwards, extends along the whole length of the bran- 
chial region. Another pair of minute simple lappets (II) is well 
seen just in front of this, and the mode of staining of the fossil 
suggests that there is a long and narrow fringe of skin (I) along 
eacli side of the rostral region. The vertebral centra are much 
constricted and sniootli, but where broken they seem to exhibit 
traces of some secondary calcification round the primiti\-e double- 
cone. Their arches are not distinguishable, but the comparative 
shortness of the centra in the front part of the caudal region 
evidently results from diplospondyly. The large pectoi'al fins are 
remarkably rounded, slightly longer than wide, and the stout 
basal cartilages do not extend more than half-way towards the 
distal margin. The long unjolnted proximal radial cartilages are 
well seen in the left pectoral. The tapering ascending parts of 
the pectoral ai'cli are crushed backwards. Tlie pelvic Hns, which 
are much longer than wide, are supported in their basal half by 
very stout radial cai-tilages ranged along the basipterygium, which 
is not produced into claspers. The individual represented is 
therefore female. Tlie two doi'sal fins are crushed downwards to 
the left side of the fossil, and seem to have been nearly equal in 
size; but the parts projecting bej'ond the edge of the tail 
probably represent only their apical halves. The first dorsal, 
which is rather fragmentary, arises just behind the end of the 
pelvic fins, while the second must have been completely in advance 
of the anal. The anal fin is relatively small, short, and rounded, 
and close to the lower lobe of the much extended caudal fin. The 
lower lobe of the latter is clearly notched near its distal end. 
The head, trunk, and fins are completely covered with very fine 
shagi'een. Most of the granides appear to be flat and smooth, 
but some are pointed, and a few on the back of the trunk between 
the pectoral fins are both pointed and slightly enlarged and 
coarselj^ fluted. 

Affinities. — So far as preserved, there is nothing in the fossil 
thus described to separate it from the existing genus Crossorhinus, 
but it is distinguished from all known species* by the simplicity 
of the dermal lappets fringing the head, and by the relatively 
lai-ge size of the pectox'al fins. 

Family PROTOSPINACID^, nov. 

Body depressed, but base of pectoral fins not produced forwards. 
Vertebral centra well calcified (pi-obably tectospondylic). Radial 
cartilages of paired fins not extending to the margin ; two dorsal 
fins on the tail, each with an anterior spine ; anal fin present. 

* Compare C. Tate Re^an, P. Z. S. 1908. pp. 354-357, pi. xi. fia;. 2, pi. xii. fig. 2 ; 
also Ann. & Mag. Nat. Hist. [8] vol. iii. (1909), p. 529. J. Douglas Ogilby & 
A. R. McCullocli, Jom-n. Roy. Soc. N. S. Wales, vol. xiii. (1908), pp. 269-280, pi. xlii., 
pi. xliii. fig. 1. For skeleton see also W. A. Haswell, Proc. Linn. Soc. N. S. Wales, 
vol. ix. (1884), pp. 92-98, pi. i. tigs. 6-8, pi. ii. fig. 13. 



TWO NEW ELASMOBRANC'H FISHES. 233 

Genus Protospinax, iiov. 

Suout short and obtusely rounded. Teeth small, compressed 
to a sliai'p edge. Pectoral fins extending as far backwards as the 
pelvic pair; dorsal fin-spines large, laterally compressed, and 
smooth, the first inserted opposite the pelvic fins ; anal fin veiy 
small, close to the elongate-ovoid caudal, which is not notched. 
Shagreen dense and fine, none enlarged ; lateral line supported 
by a series of calcified ringlets. 

Protospinax annectans, sp. n. (PI. I. figs. 2, 3.) 

Specijio Characters. — -Attaining a length of about a metre. 
Length of cranium slightly less than one-fifth, length of caudal 
fin about one-sixth of tlie total length. Teeth smooth and 
lozenge-shaped, their sharp-edged crown sometimes with a 
prominent middle point. Antero-posterior measurement of 
pectoral fin about equal to the length of the cranium, and nearly 
twice as great as the length of the pelvic basipteiygium. Dorsal 
fins about equal in size, the first arising slightly in advance of the 
middle of the fish. 

Description of Type Specimen. — The fossil, which is shown of 
one-sixth the natural size in PL 1, fig. 2, is very fragmentary, 
but there are definite points of contact between the pieces of 
rock in which it is contained, and most of it is preserved in 
counterpart, so that its general shape and proportions ai'e recog- 
nisable. The head and trunk are seen directly from above, 
while the greater part of the tail is exposed in side-view. The 
edges of the head and fins are sharply outlined by fine dense 
shagreen, while the distinctness of part of the margin of the 
caudal region is due to fossilised muscle. The cranium is well 
calcified in the usual small tesserse, and evidently not much 
distorted. Its postorbital part is about as broad as long, and the 
postorbital processes are small and slender. There is very little 
constriction between the orbits, which are completely within the 
hinder half of the cranium. The olfactory capsules form relatively 
large rounded latex'al prominences in the middle of the cranium. 
The rostral part is short and wide, not tapering but nearly 
truncated in front, and remarkable for the large size of the 
elongated anterior fontanelle which extends backwards between 
the olfactory capsules. There is no indication of a posterior 
fontanelle in the cranial roof. The jaws are not seen, but there 
appears to be a vague trace of the mandibular articulation on the 
right side well behind the occiput. All the vertebrae are crushed 
and broken, but they show much secondary calcification round 
the primaiy double-cone, and this seems to have been in concentric 
laminae (on the tectospondylic plan). As in many other fossil 
Elasmobranchs from the lithographic stone, the body-muscles are 
well preserved ; and it is clear that while in the abdominal region 
each myotome corresponds with one vertebral centrum, in the 
anterior part of the caudal region each myotome comprises two 



234 DR. A. SMITH WOODWARD ON 

vertebral centra. Thei^e is thus the common diplospondyly. Tlie 
vertebra,! arches ai^e scarcely seen, except witliin the caudal fin, 
which is displayed in direct side-view. Here the hpemals are 
apparently stouter and less inclined backwards than the nenrals. 
The pectoral arch is only imperfectly shown, but the right 
pectoral fin is complete. It is relatively large, and the supporting 
cartilages extend only about half-way from its insertion towards 
the digtal margin. The three basals are distinct, the propterygium 
being comparatively small and narrow, the triangular meso- 
pterygium about as wide as long, and the metapterygium longer 
than wide but very little produced backwards. The radial 
cartilages, which are not much longer than the basals, are rather 
sparsely arranged and do not clearly exhibit any transverse 
articulations. About 12 are arranged along both the mesoptery- 
gium and the metapterygium. Faint striations are seen in part 
of the fin-membrane, but there are no distinct remains of dermal 
i-ays. The pectoral fins extend as far backwards as the pelvic 
fins, which are much smaller. The pelvic basipterygium is long and 
gradually tapering, and bears at least 17 radial cartilages, which 
(like those of the pectoral) are not closely pressed together and do 
not show any transverse articulations. The cartilages occupy only 
half of the total expanse of the fin. As they are imperfect behind, 
the sex of the individual is uncertain. Of the median fins, one large 
dorsal, bordered in front by the remains of a smooth, laterally 
compressed spine (cZ'), arises just behind the origin of the pelvic 
fins; but it is very imperfectly preserved. The impression of the 
fin-membrane shows some fine striations, which may perhaps 
denote strengthening dermal rays. As the tail of the fish is 
relatively long, this is doubtless the first dorsal fin, but the fossil 
is too fragmentary to exhibit the second dorsal. The caudal fin, 
which is displayed in direct side-view and only incomplete at the 
upper extremity, is long and ovoid, with the lower lobe the laiger. 
The membrane here again shows faintly some fine striation. Just 
in advance of its lower lobe, a small deep and narrow fin is shown 
(«.), with most of the outline defined by oxide of manganese. 
Though its separation from the caudal is a little obscured by the 
rough fracture of the rock, it is almost certainly distinct and may 
be regarded as an anal fin. Fine shagreen covers the whole of 
the ti'unk and fins. Near the margins it is especially smooth and 
dense, but on parts of the trunk the granules are rather stellate. 
On the trunk in front of the pectoral fins, and again on the 
tail just behind the pelvic fins, it is interesting to notice that the 
coui-se of the lateral line is marked by a close series of incomplete 
ringlets (fig. 2a), as in Chimajroids and in the extinct dog-fish, 
Mesiteia. 

Young Specimen.- — A second specimen in the British Museum 
(No. 37014), from the same formation and locality, only 30 cm. 
in length, evidently represents a young individual of the same 
species. ' The cranium and vertebral column are in undisturbed 
series, with the two dorsal fin-spines in their natural position on 



TWO NEW ELASMOBRANCH FISHES. 235 

the tail ; hut the parts of the paired fins are scattered, and only 
fragments remain. The specimen, however, is of special impor- 
ta,nce, because displaced portions of both jaws with groups of the 
teeth are also preserved. The characteristic large anterior fon- 
tanelle in the cranium is well displayed as in the type specimen. 
The teeth (fig. 3) are relatively small and closely arranged, several 
series evidently functional at one time. Their exact shape is 
difficult to determine, but they seem to be transversely elongated 
rhomboids, with a, low crown, which is siTiooth, compressed antero- 
posteriorly to a sharp edge, and sometimes rising in the middle to 
a little cusp. Many of the vertebra,l centra clearly exhibit the 
secondary calcification round the primary double-cone. A frag- 
ment of a pelvic fin seems to denote a male individual, and a row 
of slightly enlarged, pointed shagreen-granules nnay have belonged 
to the clasper. The two dorsal fin-spines (fig. 3a, d}, cF), though 
fractured, are shown to be nearly similar in size and shape, and 
their length equals about one-quarter of the distance between 
their insertions. The spine of the first dorsal is supported not 
only by a short triangular cartilage behind, but also by a larger 
and more extended cartilage in front. Traces of the fine stellate 
shagreen are seen on the rostrum. 

Affinities. — The new genus and species now described evidently 
represent a family closely related to the Spinacidre, but still re- 
taining the anal Hn and a less specialised dentition. Protospinax 
is indeed a generalised type such as might be expected among 
Jurassic E^asmobranchs when the Batoids were beginning to be 
differentiated. The Batoids themselves were first represented by 
the Rhinobatidfe. and it is interesting to notice that one member 
of this family (Belemnohatis) v.oi\te.Tsi\>oYi\.Yj with Protospinax had 
a spine in front of eacli of its two dorsal fins. 



EXPLANATION OF PLATE L 

Fig. 1. Cros.^orliinns Juraasicvs, sp. 'n. ; nearly complt'te fish, nat. size. — Litho- 
graphic Stone; EichstSdt, Bavaria. I, 11,111. the three paired dermal 
lappets. . [British Museum no. P. 11211.] 

2. Protospinax annectans, gen. et sp. n. ; fragmentary fish, ono-sixtli nat. 

size. — Ibid. a. anal fiu ; d^. spine of first dorsal fin. [British Museum 
no. P. 8775.] 
2a. Ditto ; portion of lateral line of same specimen enlarged four times to show 
supporting ringlets. 

3. Ditto ; group of teeth enlarged ten times. — -Ibid. [British Museum 

no. 37014.] 
3a. Ditto; portion of tail pf same specimen, showing dorsal fin-spines ((V-, d~), 
nat. size. 



ON PATHOLOGICAL STATES IN EVOLUTION, 237 

14. The Function o£ Pathological States in Evolution. 
By MoitLEY Roberts*. 

[Received May 7, 1918 : Read June 11, 1918.] 

That dissatisfaction with much orthodox biological opinion is 
growing can hardly be denied. Not a little of this feeling is 
due to the fact that what is often given as explanation cannot 
be resolved into factors capable of appreciation, and, possibly, of 
measurement, by the intellect. The theory has to be accepted 
as more or less a matter of faith. Where there is a general 
tendency to rely on authority, speculation is discouraged, for 
oi'thodoxy everywhere rests on the native conservatism of man, 
and even the revolutionary is capable at last of fatigue. As a 
result, tentative hypotheses offered by the great leaders tend to 
become objects of faith, and among their less enterprising 
followers there arises a more or less fervent conviction that, 
however unsatisfactory they appear now, they will presently 
become demonsti'ation. Thus the theory of the germ-plasm, 
even in its later modified form, seems held too dogmatically by 
many: the 'nature' of inherited living iliatter accounts for 
every organ as it appears ; while all changes are due to obscure 
variations of an advantageous kind which give the survivors in 
the struggle a better chance. On analysis, such opinions do not 
seem truly scientific, for the " nature " of the germ-plasm can 
barely be distinguished from the directing entelechy of Driesch, 
and if the Weismannian cloud of ids and biophors is now 
somewhat condensed, the magic determinant still remains in a 
concealed vitalism which is exactly analogous, as regards the 
organism, with pantheism as regards .the universe. Nor, if we 
are told with certainty that altered characteristics are not trans- 
mitted, is the theory of small advantageous variations much more 
satisfactory, if we know neither how they come, nor how they 
are inherited. To say so much must not be regarded as treating 
with disrespect its gi'eat author, without whom we might still be 
wandering in the barren field of teleology. 

To regard these theories as hasty and, perhaps, unsound 
explanations is not to accept without scrutiny the theory 
of the transmission of acquired, or modified, characteristics. 
Though this is a view that can be defended on the physico- 
chemical grounds of catalysts which are measurable determinants 
of a really scientific order, experiments to pi'ove the fact must 
take a very long time, and we are compelled to rely on other 
methods of pi-oof. That the experiments of Tower and Kam- 
merer, for instance, suggest the transmission of modifications 
cannot be denied. Such as oppose the general view that the 
environment has thus an inheritable moulding influence on the 
organism, seem to reply that these are only rare and doubtful 

* CommunicRted by the Seceetjlrt. 



238 MR. MORLEY ROBERTS ON THE FUNCTION OF 

cases, whereas the theory of inherited advantageous variation?, 
whether continuous or discontinuous, can be made responsible 
for the whole of the phenomena. As the conclusion is gradually- 
being strengthened that large variations of a Mendelian character 
deal with other characteristics than those which are racial, all 
who rely on inherited spontaneous variations are forced back on 
the Darwinian view that small variations can gradually, if of an 
advantageous kind, convert one species into two or more, and 
that all living cliaracteristics, or oi^gans themselves, are dvie to 
such a cumulative effect. It is, of course, inferi-ed and definitely 
tStated by D irwin, that any variation in the least degree injurious 
would inevitably be destroyed. It is this statement I propose to 
examine, and for the purpose of such an enquiiy it must be 
clearly undei^stood what is meant by the word ' disadvantageous ' 
or injurious. 

At first sight nothing seems clearer. Why should we doubt 
that any functional or organic failure is a handicap in the 
biological race? By functional trouble of which the cause is 
not obvious ■ we mean some hindrance, which may be recovered 
from, to normal or physiological action. It is due to factors 
which, for the most part, are unknown. "We do not doubt that 
there is a failure somewhere, which, as regards certain cells, 
might be called organic, but often we cannot do more than guess 
where the actual failure occurs. In that advanced disorder of 
function which has visible lesions and destruction or irremediable 
alteration of the individual parts of the machine there is un- 
doubted organic disease. Can anything seem more certain than 
the conclusion that any organism which fails in the established 
functions of its species is as a fact severely handicapped, that 
the variation is disadvantageous and cannot possibly be trans- 
mitted either directly or by survival ? There are, however, some 
reasons for believing that this inference is inaccurate and that 
the function of disease in evolution is of much greater import- 
ance than that of mere elimination. But pathology has very 
naturally been neglected as a study by biologists. On the views 
generally held, it has seemed sufficient to recognize that disease 
destroyed organisms which obviously left ofispring, if it left 
them at all, that were handicapped even more heavily than their 
parents. It has been understood that their elimination was only 
a matter of time and that neither their virtues nor their failures 
eould influence the race. 

If there is one thing more than another which has struck me 
when attempting to study these questions, it is that too many 
men of science appear to believe that any serious investigation 
of other branches than their own is for them a waste of time. 
The physiologist ignores the pathologist, who in his turn is far 
too likely to fix his eyes on morbid phenomena which cannot be 
properly appi-eciated save by those with a knowledge not only 
of normal function but of the genex-al physiology which underlies 
jt. The same can be said of most workers in science, but in no 



PATHOLOGICAL STATES IN EVOLUTION. 239 

case is it more likely to occur than in that of the biologist, who, 
by the very name and nature of his task, should include in his 
apparatus a considerable knowledge of everything which deals 
with the organic, and even inorganic, world. Science, however, 
is kept in more or less water-tight compartments, and it seems 
left to the mathematician to hold the opinion that his own 
branch of learning has, somehow or another, deep relations with 
all things, including life itself. Even by him it does not seem 
to have been pointed out that in things living and non-living 
certain principles of construction rule alike. However mvich 
they were wedded to mechanico-physical explanations, biologists 
have assuredly often ignored the fact that any organism is con- 
struction, and knowing little of the laws of consti-uction have 
ignored basal facts familiar to every architect or even every 
artisan. It was reserved for Wolft', in formulating his law of 
bone-growth and reaction to stress, to propound a principle more 
far-reaching than he recognized, when he showed that living 
bone, reacting to normal or abnormal stimulation, can be proved 
to' develop in accordance with the principles of engineering and 
architecture. This law may, I feel assured, be extended to every 
living tissue, and in such an extension will be found the key to 
many phenomena still awaiting explanation. 

To one who holds this view, the work lately done by Starling 
on the " Law, of the Heart," which shows that the force Avith 
which the heart contracts is directly proportional to the length of 
the muscular fibres at the end of the preceding diastole, is by 
no means surprising. It is indeed on a par with the conclusions 
of Wolff as regards bone, and might, I believe, have been deduced 
from it or from the form I suggest, provided it is understood 
that each varying tissue has its own acquired typical reaction. 

If, then, it can be shown that disease has had a profound effect 
upon the evolution of all organisms, and that analogous results 
are found in every kind of human constructive effort in such 
numbers as to suggest as a law that all great variational develop- 
ments result not from the happy-go-lucky aggregation of small 
advantageous variation or from discontinuous variation, whether 
of a Mendelian character or not, but rather from partial failure 
and repair, we seem to be in sight of a general principle of pro- 
found importance. If this principle proves sound, it is obvious 
that immense labour has been spent by biologists endeavouring 
to explain life without seeking help from other workers. Though 
they may show some general knowledge of the cell, and even 
special knowledge of the reproductive cells, I find few who appear 
to have studied general embryology, to speak only of one branch 
of physiology. On the other hand, many physiologists and patho- 
logists have done good woi'k in some branches of evolutionary 
theory. Blan<l-Sutton, in his fruitful little book 'Evolution aiid 
Disease,' pointed out that "Pathology is only a depai^tment of 
Biology, a.nd it is important to bear this in mind in studying 
disease." It is true that he went little further than to show that 



240 MR. MORLEY ROBERTS ON THE FUNCTION OP 

what is pathological in one organism may be physiological in 
another and that many diseases are reversions, that is, failure in 
normal growth. Yet this greatly needed to be shown, and it 
is not to be expected of a great pathologist and sm'geon, and 
perhaps the less the greater he is in his own branches of work, 
that he should attempt tasks from which many of the biologists 
themselves seemed to shrink. Claude Bernard made similar 
remarks as to pathology. It is to be regretted that a stumbling 
block was placed in the path of pi'ogress by Darwin's hopeless 
dictum as to the explanation of variation, just as another was by 
Huxley when he declared consciousness an insoluble problem. 
In every science great discoverers have too often delayed progress 
as much by authoritative unsound opinion as they have advanced 
it. Every Bible is first a book of revolution and then a lefuge 
for reaction. Yet no man can possibly know all he should know 
for the purposes of his own work. This fact aftbrds the only 
justification for those, who cannot pretend to profound knowledge 
in any special line, attempting to solve problems which by their 
nature are beyond the specialist. They may have been able to 
grasp in a measure the general conclusions of each science and 
by a happy, perhaps accidental, combination, show at least part 
of the forest to those more particularly occupied with the trees 
themselves or the floia of the undergroAvth. 

It is remarkable that hithei'to no one seems to have made the 
observation that reaction to an actual, or threatened, breakdown 
is one of the basal laws of all construction and organization. 
Yet none can read engineering without observing that -all 
development has followed such lines. As new stresses are 
introduced failure is threatened and steps are taken to obviate 
disaster. What is a patch in one engine becomes organic in the 
next. Since waste of energy can be looked on as pathological, 
we observe the reaction in the engineer against such failures, as 
the atmospheric engine is succeeded by improved forms ending 
in the quadruple expansion engine. Many other instances could 
be adduced in general or special engineering evolution, but 
the best illustration of the facts which need elucidation can 
perhaps be found in Gothic architecture. If such a demon- 
stration of this general principle can be made it will go far to 
obviate the objection, very likely to be made, that what occurs 
in human construction has no relevance to the living organism, 
especially if it can be suggested forcibly that human intelligence 
is in itself a reaction, and that the law obtains in developments 
of all kinds. That tiial and error are at the base of evolution is 
indeed implied in the current teaching as to variation, and its 
extension to intellectual processes will surprise no worker who 
has had to deal experimentally with the unknown. We may 
e:3^ect, but never know where to look for, failure till we see it. 
When it is seen we can do our best, as reacting agents, to remedy 
it. Having said so much, and leaving aside the wider implica- 
tions of such views, we may turn to such a problem of construction 



- PATHOLOGICAL STATES IN EVOLUTION, 241 

as tlie evolution of a cathedral, in the hope that it may throw a 
light on other than architectural puzzles : merely observing, on 
the way, that no general principle yet discovered is confined in 
its application to one Vjranch of knowledge. Having once found 
it, our task is to employ it as a weapon of further analysis. 

It is more or less a commonplace that function creates 
structure, however Lamarekian that may sound, and in the case 
of architecture of a religions order the function which constructs 
is public worship. Tn hne climates the necessary structure is 
often a rooHess temple. In tropical climates a Hat roof may be 
needed as a protection against the sun. In temperate climates a 
walled enclosure is insufficient and a |3.at roofed structure cannot 
keep out rain effectually or bear heavy snow. Thus arose the 
pointed or sloping roof. Rut it has been said that " Gothic 
architecture is not a style. It is a fight." The ai'ch is a mighty 
warrior. It gives and receives thrusts. The slojjing roof partakes 
of the same nature. Need created it and the nature of materials 
and the positional energy we call gravity caused thrusts which 
endangered the simple walls of the building, walls at first meant 
to endure nothing but flat roofs probably covered with brush or 
the like material. To build stronger walls might have occurred 
to the primitive architect, but as the danger was immediate, he 
probably at once shored those in existence, and then built others 
at a right angle to act as buttresses. In the meantime the 
worshippers increased in numbers, and it is indulging in no 
flight of fancy to suppose the later builder saw that if the new 
external walls were roofed over and doorways cut into the main 
building, there would be an immediate increase of space by the 
creation of chapels. Such a series of embryonic additional walled 
spaces, with further door\^'ays in them leading to each other, 
obviously gave him the aisles. The flying buttresses which are 
such a feature in great Gothic architecture had, I can only 
suppose, a like origin. They were originally buttress walls carried 
up to the roof. At some period a genius, already acquainted 
with arcuated structure, saw that if the inside of these walls was 
cut away they would still take a heavy thrust and lighten the 
rest of the building. If, however, on being converted into such 
slender stone shores they showed signs of yielding, what could be 
easier than to pile some of the material taken away on the base 
of the flying arch and thus create the beginning of the pinnacle ? 
Though an architect might develop such a rough statement, he 
would be the first to admit that it represents in few words much 
of the evolution of a church; that is, he would own the structure 
sprang from need, and that each new need caused a constructional 
failure which, when strengthened and corrected, was the cause of 
further structure. He would further tell us that all good orna- 
ment is organic ; that it springs naturally from the work already 
done, being in its origin just the little more needed to give a 
margin of safety, though on it later are exei'cised the iBsthetio 
faculties of man, which ai'e again a response to the need of full 



242 MR. MO RLE Y ROBERTS ON TUE FCNCTION OF 

satisfaction for the instinct of workmanship. Human ornament 
is in fact strongly iiomologous, if we may use that word here, 
witli the beauty of very energetic birds, wlio cairy out by virtue 
of their free energy the extension of sti-uctures and colours 
already existing in their less brilliant forms. That, however, is 
by the way. The main fact we are concerned with is that tlie 
structure as a whole evolved through trial and error, through 
failure and repair, through a threatened structure to a moi'B 
complete and adequate one for increased function. In a woi-d, 
the great origin of structure was failure after failure duly com- 
pensated for. Is there any reason for believing that variation 
in the structure of living ^organisms follows exactly the same 
principle 'i Are we entitled to say that the mammal, for in- 
stance, with all its complexity, is the result of infinite ages of 
functional failure or disease which was met by processes of i-epair 
and reaction ? In a word, can we speak of the evolutionary 
value of disease, of impaired function, of disadvantageous varia- 
tions 'i It seems possible to do so, if what is ti-ue of one structure 
is roughly true of another. 

It may seem absurd to speak of the value of disadvantageous 
variation, but it is no more absurd than to imply that all 
vai'iation is advantageous because it is perjietuated. What is 
useful at one period may be harmful at another, and embryolo- 
gists thoroughly understand that developments useful in fcetal 
oi' larval life may open up many dangei'S for the adult, Tlie 
real point to be considered is whether organisms as species do 
not vary and run great, even largely destructive, risks by an 
increased pressure of function which, in the few that finally 
react or whose descendants react to such stress, results at last 
in structure that is advantageous as cdtered. The given variation 
in itself may be a failure of what was normal function in the 
species, and we should therefore as pathologists or physiologists 
speak of it as disease, but if the few that recover become a new 
species, a mended race, it is no longer disease. After many 
generations it may be truly advantageous to individuals. Have 
such processes occurred in the evolution of organisms as they 
undoubtedl}^ have in the arts and social progress, where we often 
observe political failure or organization result in ad hoc I'eaction 
which leads to a changed social form? I have no doubt that 
they do, and many organs in mammals, to speak only of them, 
show it. It is, in fact, a universal principle. As beavers patch 
up a dam when it yields or threatens to give way, so tissues, 
organs, and societies react to threatened disaster. In no tissue 
is this clearer, than in bone. It is true that Wolff's law only 
deals directly with mechanical stresses, since it runs — " every 
changa in the form and position of the bones or their function is 
accompanied by certain definite changes in their internal archi- 
tecture and by equally definite secondary alterations of their 
external conformation in accordance with mathematical law;" 
but I hope to show reasons for concluding that such a law may 



PATHOLOGICAL STATES IN EVOLUTION. 243 

be stated in more general terms and applied to every tissue and 
organ, provided we add that the more complex the tissue or the 
organ the greater the liability of failure, and that each tissue 
reacts in a typical way. 

It is unnecessary to go into details of osteogenesis and mor- 
phology. It has been recognized by engineers that the head of 
the femur is formed exactly in accordance with mechanical law. 
Had any of them been required to design a structure fit for 
undergoing the stresses borne by the femur in its development 
and after-life, he would have sketched a figure extremely like it, 
not only in its general shape, but in the trabeculee which suppoi't 
the bone in every direction where extra stresses are applied by 
normal function. The important point to note is the fact that 
femoral developement follows stress in individual development, 
from which we must draw the conclusion that it followed stress 
during evolution, not that its value for complex function was 
gradually increased by chance or "spontaneous" variation, unless 
we attribute to "spontaneous" a meaning which Darwin never 
gave it, seeing that he denied knowing how variation arose. All 
the variations were definite responses, and it is easy to infer that 
before response became rapid and easy every kind of disaster 
and disablement must have occurred to those subjected to re- 
action-provoking stresses. The very process of adaptation (and 
on these lines "adaptation" is no longer a mystic word) implies 
long periods of disordered function and poor structural response 
even in those who survived after repair. But now bone is so 
plastic and fluent that when it is grafted the osteoblasts and 
osteoclasts shape it according to the form of the main bone of 
wliich it becomes a part. 

When we speak of repair it may be noted that the treatises 
on this subject are strictly limited in their pui'view. They mostly 
follow Hunter, a vitally important figure in the history of 
pathology and indeed of all medical science, who, however, 
lacked the apparatus of knowledge now at every one's disposal. 
We learn a great deal about the repair of wounds and fractures : 
of the functions of the fibroblasts or of the wandeiing cells 
of the blood-stream, and are told, lately, much of regeneration, 
but of the evolutionary value of organized exudations we hear 
nothing. ISTor has it been suggested that it is to this and 
analogous processes that mvich new structure is due. That this 
is so is strikingly apparent, as I shall attempt to show, in many 
organs of a highly specialized type. In no structure, perhaps, is 
the process so clearly seen as in the mammalian heart, which is 
a perfect museum of evolutionary failures and dislocations, com- 
pensated for by an extraordinary complication of patched-up 
tissues and organized exudations in which, perhaps, one tissue 
takes on the functions of another and some evolutionary rem- 
nants long survive without function. I was, indeed, first led to 
take this genei-al view of the variational value of pathological 
conditions by observing that the heart, when laid open from any 



244 MR. MORLEY ROBERTS ON TUE FUNCTION OF 

aspect, powerfully suggested an organized or cured aneurism. 
Many must have made the same observation, even if they have 
not come to similar conclusions. The anatomist and pathologist 
perhaps know their subjects too well and are necessarily greatly 
dominated by current theory. The general adaptation of the 
heart to the work it performs may well delight the anatomist as 
he studies its machinery. His main business is not evolution. 
The pathologist on the other hand, observing its many failures, 
is scarcely likely to discern that by failure itself may come 
eventual perfection, and while the physiologist considers its 
functions rather than its apparatus, he studies it as it is, not as 
it was. In each case the observer may not see the forest for the 
trees. Yet when we look at the partially repaired aneurism with 
its fibrous growths and turn to the opened heart, the essential 
relationship of the chordca tendinece, for all their definite functions, 
to the rude fibres of an aneurism is obvious. Is such a likeness 
an accident of evolution and pathology, or are we to consider the 
heart as much an organized dilatation sac of the whole fused 
circulatory canal as the cured aneuiism is of a. part of it ? It is 
in embryology that we seek for confirmation of what is suggested 
by anatomy. But even anatomy alone offers powerful proof of 
the view that the heart, as we know it, is the latest result of 
i-epeated failures of the circulatory canal under strain and of the 
repairs effected by the stressed tissues in their response to 
changed and abnormal stimuli, just as bone alters under its 
particular stresses. During .embryological life there is found in 
the heart a small patch of non-functioning muscle in the anterior 
segment of the mitral valve. Its presence is intelligible if we 
consider it a relic of a disrupted and repaired organ. The 
muscles of the heart are obviously homologous with those of 
the arteries. Yet they have become striated although they are, 
of course, still involuntary. IS on-striated muscle is the earliest 
in evokition. It seems that the increased functioning of the 
cardiac muscle has converted it into its striated form so that it 
resembles skeletal muscles, which are much more active than 
non-striated muscle. The whole histology of cardiac muscle 
probably represents the result of great stiains. Structures such 
as the disks or bands of Ebarth are found no whei^e else and may 
be the result of peculiar stress. There are even portions of 
muscle which no longer perform muscular functions. Their 
fibres do not contract but serve instead to conduct stimuli as 
if they were nervous tissue. All tissue is conductive, but the 
bundle of His, with its Purkinje fibres, which carries the impulse 
from the sino-auricular or Keith-Flack node to the venti-icle, 
transmits messages at ten or twelve times the normal muscular 
rate. When it fails there is heart block. In the embryo the 
valves arise from the cardiac walls and are composed of muscular 
tissue which by the action of fibroblasts gradually become non- 
muscular. This must have been originally a pathological process. 
It is a reversion, a degeneration made use of. We observe 



• PATHOLOGICAL STATES IN EVOLUTION. 245 

analogous, or shall I say homologous, results in the hypertrophied 
heart. The normal xnale heart weighs about eleven ounces. In 
some cases of aortic stenosis it may weigh over thirty ounces. 
In such hypertrophied muscle are often found fibrous tissues 
which probably represent the connective tissue of muscular fibres 
Avhich have atrophied from overstrain. The chordoi tendinece of 
the mitral valve are less muscular and more fibrous than the 
same attachments of the tricuspid. This adaptation difference 
lessens strain on the thinner right ventricle. It has, indeed, 
i-emained thinner on that account. In the reptile with a function- 
ing foramen the valves are purely mechanical, as pressure is 
relieved by the patent orifice. The fossa avails in the mammal 
is a remnant of the early communication between the auricles. 
In a large number of normal hearts there is a small valvular 
passage yet remaining in the left margin of the fossa. None of 
these phenomena seem capable of explanation as the result of 
spontaneous variations arising from some theoretic instability of 
the organism. To ai-gue that they are is to give biologic mystics 
a chance. It appears obvious from all these facts taken together 
that cardiac evolution has been a series of caused variations due 
to increased and varying stresses which acted not only as' a 
moulding force on the shape and musculature of the heart but 
on all its appendages. In the muscle of the ventricular walls 
with its extraordinaiy complexity of layers and interlaced fibres 
lies powerful evidence of such reactions. In both ventricles 
there are seven muscular layers, while in the arteries there 
seems but one. In the left ventricle these layers are obviously 
thicker and stronger than in the less stressed right cavity. But 
how did the ventricular cavities acquire more layers than the 
arteries? There is obvious reason for believing that stress can 
be responded to by increase of muscle fibre during evolution. 
In the gravid uterus the smooth fibres of the wall inci'ease to 
eleven times their normal lenglh and are from two to five times 
as broad. There may be new fibres in it. I doubt if any one 
knows. But in evolution new fibres are undoubtedly found. In 
the arteries, the fibres of non-striated muscle in the tunica media 
are for the most part circular, but they appear to have more or 
less longitudinal branches which interlock with like branches of 
the neighbouring fibres. One of the most prominent features of 
an individual aneurism is the thinning out, and sometimes the 
disappearance, of the tunica media. The muscle fibres in such 
cases are completely broken down, and if the aneurism is repaired 
in individuals the work is done mostly hy an increase of the 
connective-tissue elements. The process is said by some to be a 
reparatory endarteritis, in which the tissues of the adventitia 
proliferate actively. But the evolutionary process has obviously 
taken the path of increase and reactive proliferation of the 
muscular elements of the media. 

It is often observed that the aneurism which displays sutHcient 
reacting power to the stresses of the blood strea,m accumulates 

Proc. ZooL. Soc— 1918, No. XVIIT. 18 



246 MR. MORLEY ROBERTS ON THE FUNCTION OF 

blood clots iu layers, and it is therefore all the more interesting 
to note that in the embryo the columnce carnece and chordce 
appear to rise from a spongy network which at an early age fills 
the primary ventricle. Such origin is strongly suggestive of 
some process analogou.s to blood clotting or to an iri-itative 
reaction of the embryonic ventricular wall. The evolutionary 
dilatation sacs which I suggest were originally pathological can 
be seen in the embryo during the process of their formation. 
The path laid down by pathology is trodder by physiology. It 
follows that during evolution there must have been an immense 
destruction of organisms whose circulating canals did not react 
and numbers which retained their unaltei'ed "specific ''characters. 
The same process goes on to-day. Though many die of cardiac 
disease, it may be that much youthful functional trouble and 
•even more serious adult disorders are even now remoulding the 
heart. No organ is perfect : if it does not degenerate it pro- 
gresses. Though such processes are " disease," it by no means 
follows that they will hQ destructive, any more than that the 
functional incapacity of the tricuspid valves in athletes, Avhich 
probably precedes what is known as "second wind," is anything 
now but a cardiac safety-valve. 

As we learn more of the heart and its latent capacities we 
may perhaps say with the late Dr. H. G. Sutton, "we trust 
nature too little, to say the least of it." But there are, of course, 
great difficulties to overcome before we can hope to understand 
how the cardiac musculature has altered and may still be 
changing by the addition of new fibres. As yet, little is known 
of myogenesis. Like a neurone, a muscle cell seems to last a 
life-time, and though both may degenerate or die, neither pro- 
liferates after the early period of development. But whatever 
their histogenesis, new fibres do appear in evolution. Harvey 
did not refuse to believe in the validity of his own conclusions 
because he lived before Leeuwenhoek. With considerable hesi- 
tation I ventui-e to suggest that morphogenetic stress is at its 
height during foetal development. The child in totero has not, 
perhaps, the calm and happy life commonly attributed to it. On 
the contrary, it probably leads a sti-enuous existence, and if it 
inherits a new weakness this is shown just where and when new 
stresses find plastic embxyonic tissues to respond to them. If 
such a speculation is sound it accounts for many phenomena. 
But in any case, whatever the machinery of inheritance and 
evolutionary repair, it is certain that new fibres arise where they 
are needed. 

If svxch views in any way represent the biological history of 
the heart, it is obvious that many of the opinions of variation 
usually held are without foundation. Every variation. is definitely 
caused ; it is in no sense accidental or spontaneous ; it may not 
be even at once advantageous to the individual : on the contrary, 
it may be a severe handicap which puts greater general stress on 



PATHOLOGICAL STATES IX EVOLUTION. 247 

all who experience it, though suoli stresses fall short of those 
which cause death. Yai-iations of this oixler may only be 
advantageous to the whole species as a continuing rpce. They 
may destroy, and doubtless have destroyed, individuals without 
number at an earlier age than the usual, life-period of the 
unvaried type. We may possibly imagine a part of humanity, 
now responding to stresses which make the heart do njore work 
and fail earlier, displaying such energy during their shorter 
life as to. displace those with a normal cardiac mechanism which 
survives to the average age of man. It is to be inferred from 
these considerations that the structure of an organism is not a 
congeries of minute fortuitous advantageous variations, nor the 
gradual massing of details in an orthogenetic line, nor the result 
of large discontinuous variations due to chromosomatic in- 
heritance, but a complex of definite I'eactions to definite stresses. 
The true theory of living structure is that its growth is neither 
casual nor foreseen, but that is what we may call, in political 
language, the " opportunism " of the organism as a whole. Every 
advance is a forced, even a desperate, experiment. Life, like a 
hypothesis or a dam, is built up by stopping leaks. 

The evolution of the stomach seems to have followed the 
lines suggested for cardiac development. From the physiological 
point of view, a straight intestinal tube which becomes dilated 
cannot be considered anytliing bat pathological. It has failed 
under the stresses imposed on it, but the organism which 
reacted turned a weak dilatation sac into a strong perma- 
nent food pouch. Tlie results to the reacting organisms were 
many. The ingested food became temporarily static, was more 
thoroughly dealt with, and the organism was not continually 
feeding. Its whole available energy was not devoted to nutrition : 
it had time at its disposal and could develop other functions 
leading to further structures. That the human stomach is such 
an organized failure is suggested forcibly by the musculature. In 
the small intestine this is composed of two layers of fibres, 
circular and longitudinal. In the stomach it is made of three 
sets, an inmost layer of oblique fibres being added. This oblique 
layer is obviously a later growth and, as would be expected on 
the lines laid down as to disaster and repair, its strongest fibres 
are found just where they are wanted, that is, supporting the 
greater curvature or dilatation of the stomach. This later 
layer is naturally less well developed than the longitudinal and 
circular fibres. Other oblique fibres are foi'med about the pylorus 
where they form the sphincter. I suggest that these oblique 
muscle fibres arose as points of strain, under intense stimulation. 
The dilated pouch has reacted in accordance with mechanical 
law, just as the heart did with its more complex arrangement of 
oblique fibres woven into a structure capable of giving in the 
left ventricle a thrust of over fifty pounds. The reacting organism 
is no fool of a mechanic either in its bones or its muscles, and 

18* 



248 MR. MOKLEY ROBERTS ON THE FUNCTION OF 

these phenomena are additional reasons for extending Wolff's 
law to all tissues. If protoplasm did not so react there would be 
no problems to solve. 

Such views on the mammalian, gastric apparatus are so 
obviously supported by the embryology of the orgau that there 
is no need to go into details beyond noticing that in the fourth 
week there comes the first dorsal bulging in the fore-gut. Of 
the curiously shaped fundus, Keith has remarked that it is in 
origin like the caecum, but I do not think it has been suggested 
that its form has possibly been moulded by the presence of the 
large air bubble so often seen in X-ray photographs. It is an 
elastic air-reaction pouch just as the whole stomach itself is a 
food-reaction pouch. It began to give way there, but the process 
was stayed. So far as I am aware, it is not provided with 
obvious oblique fibres. Further investigation may find them. 

If evolution is still proceeding, is it absurd to suggest that 
the common disorder known as dilated stomach may be a patho- 
logical process actually in the process of becoming physiological ? 
According to some phj'sicians, few modern stomachs do not 
suffer at times from an amount of dilatation which is patho- 
logical ; i. e., their gastric musculature fails to react correctly. 
The stomach ma}^ yet be such a functioning dilatation pouch as 
to enable the human race to do with no more than one meal a 
day or even less. Our descendants will have all the more time 
for work. This by no means implies that the empty stomach 
should be any larger than it is now in healthy subjects. Befoi'e 
the invention of X-rays the gastric apparatus was always 
pictured in text-books as usually seen on the post-mortem table. 
The dead stomach was shown as the portrait of the live one : 
the weakened pouch of the sick man as that of the live and 
healthy subject. But nowadays it is known that such extreme 
dilatation is natural only when a lai-ge meal has been taken. 
When the healthy stomach is empty it contracts so that it 
nearly resumes its ancient cylindrical character and is of a size 
not much greater than that of the small intestine. With further 
development it might hold still more and yet react in the same 
way. The suggestion that functional failure or disease which 
becomes organic and destructive in many, may, in reacting and 
surviving organisms, alter their outlook on life and all their 
activities, seems to me powerfully reinforced by these considera- 
tions. The disadvantageous variation does work, and finally 
improves the race. 

It can even be shown that disadvantageous variations actually 
become permanent racial characters. We may consider hernias. 
In the prone position of most animals, hernial sacs, now known 
not to be essentially pathological until they are forced open by 
mechanical stresses or relaxed by organic weakness, are not a 
great source of danger. They may even be considered as an 
additional means of securing the peritoneum to its connective 
tissue. During the processes of evolution, however, a mammalian 



PATHOLOGICAL STATES IN EVOLUTION. 249 

hernia seems to have occurred ahnost universally and to have 
established itself as noroially physiological. The tunica vaginalis 
propria of the testis is actually part of the original peritoneal sac, 
as can be seen in the embryo. During foetal life it is separated 
from the parent tissue. In whatever sense we now call such a 
change physiological it seems impossible to regard it as originally 
anything but pathological. Is it too startling to declare that it 
is an evolutionaiy sloughed tissue such as is often seen in 
strangulated hernias ? I certainly do not know how we can 
describe the scrotum as anything else than the coverings of an 
evolutionaiy hernial sac which is not only of no advantage but a 
positive danger to most male animals. In some, the pigs for 
instance, the testicles do not descend into an external pouch but 
are supported and protected by the normal skin tissues, not by a 
thinned and delicate integument of later development like the 
scrotum, a tissue still scantily supplied with the non-striated 
muscular fibres which might have reinforced it and are perhaps" 
now developing slowly. When we consider the i^arity of mus- 
cular fibres in human skin tissues in compai-ison with those of 
animals, their greater frequency in the scrotum and perinseum 
suggests that they are a reaction product. They act in the 
dartos, or deeper layers of the scrotal dermis, at right angles to 
the rugfe and are something of a support. The pink colour of 
this structure is due to the presence of these muscular fibres. 
They are not connected in any way with the cremaster muscle 
and therefore not afiected by the cremasteric I'eflex. In no sense 
can the descent of the testes be called a.dvanta.geous. It causes 
a weak spot, recognized as such by men and animals. The 
Japanese wrestlers are trained from youth to return the testes 
into the inguinal -canals. If the translation of the testis from 
a safe place to an exposed one has had any good results they 
have been indirect and only discoverable, though not yet 
discovered, over long periods during which the change must 
have been disastrous to many. To argue that they were advan- 
tageous to begin with is to destroy the authority of reason. 

It may seem an undue extension 'of the view that pathology 
has played an immense part in evolution if it is suggested that 
it was upoii pathological conditions that the very existence of 
the Metazoa depended. There can be no doubt that they origi- 
nated from some protozoon by a failure of normal physiological 
fission. We see here how theories of disease may be modified 
according to the point of view ta,ken From that, shall I say, of 
a protozoan Hippocrates or Hunter nothing can be more obvious 
than that a failure of mitosis would be a calamity, the birth of a 
monster, of Siamese twins, among the normally constituted uni- 
cellular organisms. It is still in the processes of reproduction 
that we find the strongest evidence of the part played by disease. 

When considering such problems in this light it seems some- 
what difficult to account for the satisfaction of many with the 
theory of small cumulative advantageous variations. What ground 



250 MR. MORLEY KOBKRTS OX THE FUNCTION OF 

is there for imagining such machinery coiihl result in a complex 
sei'ies of adaptations such as the uterus and what we may call its 
habits and customs in dealing with the embryo from the entrance 
of the ovum till birth ? Even- those who adapt to their o^^•n 
ideas some theory of large discontinuous variation will, in the 
end, be compelled to attribute the uterine giowth and functions 
to a mystic power or virtue in the original germ. They may 
follow some philosophers and " unpack "' powers out of a con- 
jurer's bag without telling us how they got there. Yet if we 
regard the uterus as the result of tissue reactions under abnormal 
stimuli, being guided in research by the processes seen every day 
in disease, the variations, whether small or large, continuous or 
discontinuous, assume an aspect neither fanciful nor mystical, 
and our need for biological faith is reduced to a decent scientific 
minimum. 

The fact that the embryo acts upon the maternal organism as 
a parasite against which the mother has to be protected, is 
commonly recognized, but I have not seen the obvious conclusion 
drawn that the whole history of the mammal must have been 
due originally to a pathological accident in some one or more of 
their ancestors. The mammalian animal still lays eggs, but they 
are not extruded. When such retention first took place, it must 
have been due to an accidental pathological delay of the travelling 
ovum, owing perhaps to catarrh of the tube. Even now the 
mother has to be rendered immune to the products of the offspring. 
Many of the phenomena of early gestation are those of immuni- 
zation, in many cases a very slow process, as is shown in human 
beings by vomiting and malaise. It has, moreover, not been 
clearly or generally recognized except by pathologists that the 
very methods by which the ovum attaches itself to the uterine 
wall a,re, so far as the hostess is concerned, actually pathological 
and bordering on the malignant. Yet they have resulted in a 
series of protective reactions which save the parent and permit 
the growth of the parasite. Tlie method by which the ovum 
becomes partially buried in the tissues is obviously of a destruc- 
tive kind and curiously analogous to the malignant processes 
seen in chorion-epithelioma. Bland-Sutton remarks, " This disease 
is instructive because the erosive action of the trophoblast is the 
physiological type of the invasiveness so characteristic of many 
varieties of cancer." It may, I think, be added that it is the 
balance established by reaction which makes the trophoblastic 
action physiological. 

That the influence of the ovum on the undeveloped tube must 
have been of an exceedinglj'^ dangerous character is now seen in 
tubal pregnancies during which the chorionic villi frequently 
penetrate the wall of the tube, which does not react as powerfully 
as the uterus. Such a process in the uterus, which is itself a tubal 
dilatation, is now normal because these villi, the earlier nutrition 
roots or organs of the parasite, are prevented from injuring the 



PATHOLOGICAL STATES IN INVOLUTION. 251 

uterine wall irrevocably by the transformation of the reactive 
uterine decidua and the chorionic villi and the allantois of the 
foetus into the combined temporary organ known as the placenta. 
It may be noted that the non-placental mammals are less 
exposed to the destructive and toxic efiects of their ofispring 
as they are born at an earlier stage than in the case of the 
deciduate mammals. The marsupial foetus is about half an inch 
in length when transferred to the milk-pouch. It is impossible 
to look at the placenta without recognizing that it is what we 
may call a compi'omise growth, one which serves the embryo 
without destroying the parent hostess. That all mammals are 
not yet fully armed against any morbid alteration of function 
in the penetrating chorionic villi is seen, as suggested above, in 
chorion-epithelioma, where the energy of the villi trophoblasts 
leads to a malignant overgrowth of the epithelial elements, 
which the maternal tissues fail to inhibit. The hydatid mole, 
which does not as a rule become malignant, is a case where such 
inhibition has been sufficient. All these cases, malignant or 
benign, support the view held by many that malignancy always 
depends on the failure of some tissue inhibition, and that if 
bacilli play any part in the drama it is that of helping to upset 
the balance between tissues which are fundamentally hostile 
though they exist normally in symbiosis. These phenomena 
regarded as a whole establish on a firm foundation the 
view that the uterus and its reactions during gestation are 
definite protective processes or variations springing originally 
from a purely pathological accident in some ancestors of the 
mammalians. However complex the embryology of the uterus 
and its appendages, the broad facts are compatible with this 
view, which is strengthened by the later parasitic history of the 
offspring after birth. The mammse appear to be a compromise 
between the needs of the infant and the protection of the 
mother : they originated in sore or tender spots on the epithelium 
most exposed to the assaults of the parasite. The growth of the 
nipple is a complex variation depending on the mechanical action 
of sucking with a reaction proliferation of the epithelial elements 
of the sweat and sebaceous glands and an increased blood-supply 
as special inaternal protections against oral infection. It seems 
to me that few stronger instances can be found of the fact that 
the development of many organs, if not all of them, is the result 
of direct reactions or adaptations which are in the nature of 
i-epair to tissues otherwise likely to suffer disastrously. 

It is large macroscopic results of this order which enable us to 
reason about other finer reactions, and even help us to link to the 
general process those of a microscopic and idtra-microscopic 
character which we class under " immunity." Such phenomena 
are reactions under stress which, by the provocation of catalysts, 
influence life. Much of human character, even, is similar re- 
action, perfect or imperfect, to the infections to which the race 



20L MH. MORLEY ROBERTS OX THE FrXCTlOX OF 

has been and still is expose*! Thus psTcliologv itself must at 
last be classed as the result of physical reactions, a conclu- 
sion fully in accord with the work of Pavloff on conditioned 
reflexes. 

If any further illustration of the conclusions so far suggested 
is necessaiy. it may be found in the growth, of the mesentery. 
It has often been pointed out that the embryonic processes bv 
which it is formed are histologically those of plastic organized 
exudations. The attachments of the whole tube do not come 
about at the same period of foetal development, and it seems of 
significanc-e when we note that the mesentery of the small gut has 
an oblique attachment, to the posterior abdominal wall from the 
duodenum to the right iliac fossa, only found in animals which 
have assumed the upright posture. This comes into existence as 
late as the foui-th or fifth month of fcetal development. Before 
this band was formed there must have been a great series of 
disasters, for even now the last part of the mesentei-y to become 
attached to the abdominal wall, that is, the angle between the 
ileum and ascending colon, sometimes remains free. A volvulus 
may easily form there by rotation of the ileo-colic loop. The 
whole history suggests a series of lymph effusions, caused bv 
pathological states, some of which were soi-ted out by the lethal 
process of natural selection, the remainder surviving and leaving 
offspring with the liability to organize the effusion in the safe 
way. The pathology of those cases in which what are known as 
Lane's Xinks can be found is obviously of a similar character. 
The stasis of the affected bowel causes lymph effusion and the 
formation of a band which is morphologic-ally homologous with 
the early mesentery. 

After reviewing phenomena such as these, the conclusion 
seems inevitable that single small favoiu^ble variations have 
not done the whole work of evolution. They may play their 
part as coiTelated changes, but they then take their place in a 
series of which the causes can be recognized. In combination 
with reasonable views of nse and disuse and of increased or 
decreased blood-supply they may, perhaps, be held to explain 
such phenomena as the delicate co-aptation of some c-ardiac 
valves. Their place in the explanation of the phenomena of 
mimiciy seems obvious. But though they may help us to com- 
prehend how tissues become finished structures if they are 
combined with the results of functional energy, they yield no 
hint as to great or decisive developments and the mechanism 
involved in them. If the reasons adduced for the thesis laid 
down carry any weight, it is obvious that many, if not most, of 
the really decisive variations in all internal structure depended 
and still depend, rot on variations which can be called favourable 
but on those that for the major portion of the organisms involved 
are directly disastrous : not on variations which are small but 
on those which are big enough to be appreciable as the cause of 



PATHOLOGICAJL STATES IX ZTOLmOX. 253 

immense functional and structural results: not on changes ^hich 
can in any sense be called sfKDn tan ecus, by which we may suppose 
is meant those no cause can be assigned to, but on variations, 
which, though thev occurred ages ago. wei-e obviously due to the 
very same causes that the pathologist can demonstrate to be 
working at the present day. Only such organisms as respond by 
'direct reactions in a manner that finally rums out to be useful, 
or at the very least compatible with life and reproduction, are 
able to survive. The whole of gi-owth and development thus 
becomes largely a function of effective morphogenetic repair to 
organic failure and disease. 

Though this is not the place to deal at length with the vexed 
question of transmission of modifications, it may be remarked 
that the foregoing arguments seem to imply that such alterations 
as a matter of fact are inherited. I think some progi-ess can be 
made if we simply assume pi-ovisionally that organisms do tend 
to repeat themselves and that it is unlOceness rather than likeness 
which requii'e? explanation. TVe know that gross unlikeness is 
almost always due to a lack or excess of some internal secretion, 
hormone, or enzvme. and fi'om this it may be infeiTed that 
likeness is due to such catalytic machinery coming over in the 
zygote, and to each differentiation producing anew its own 
peculiar products which stimulate or inhibit further git)wth and 
differentiation. Some time ago I was sti-uck by a remai-k of 
Starling's that each new oi'ganism seemed a fi-esh •• creation." 
He gave this up on account of the difficulty he found in the 
'• time element " of the problem, but I ventiu-e to think he was 
right in his surmise. There is a growing body of opinion in 
support of this view, as the names of Cunningham. McBride, 
Dendv, and BoiuTie seem to bear witness. Ve must certainly 
take into account these regulators of metabolism, and if we 
accept the view that hypo-thvroidism determines ci-etinism. or, 
in the adult, mvxcedema : that hyper- thyroidism is the' direct 
cause of the phenomena seen in Gi-aves' disease, just as h\'per- 
or hypo-pituitarism causes giantism or infantilism in children 
■while a later oversrrowth of the gland causes aci~omegftly. I see 
no difficulty in accepting the hypothesis that gix^wth is deter- 
mined. ?. e. stimulated or finally inhibited, by non-living catalysts 
or secretions not necessarily confined to the endocrine organs. 
In this way a bridge may perhaps l;e btiilt between the oithodox 
Weismannian and the Lamarckian. Growth and character are 
caused by determinants, but these are not part of the cytopljism 
itself, they are the machinery by and through which living 
matter acts. The organism is not built up by special protoplasm 
or by entelechies or by any mysterious elan crt'atif. It arises 
from the definite influence of definite catalysts originating, in an 
orderly sequence, as the organs become differentiated, while tie 
individual is as a whole esposed in an infinite progression to 
the internal and external stimuli of a like but slowly changing 



254 ox PATHOLOGICAL STATES IN EVOLUTION. 

environment to which it reacts. The factors which did the 
work are working now. 

To recapitulate the tentative conclusions arrived at, it may be 
Suggested that — 

1. Mechanical reaction to stress is a general law of all tissues. 

2. Morbid conditions in many cases give rise to repair which 
becomes physiological. 

3. Such repairs lead to new functions, new stresses, further 
morbid states and further repair. 

4. These factors are some of the main causes of specific and 
generic difierences. 

5. In all probability transmission of changes caused in the 
way indicated does take place by a morphogenetic reply in utero 
to increased fvinctional stresses. 

6. As it is a narrow view to assume that pathology in all cases 
tends to death, the study of pathology and general physiology 
should be part of the preparation of the biologist. 



XOTES ON THE BEAVERS AT LEOXARDSLEE. 255 

15. Notes on the Beavers at Leonardslee, 1916-1918. 
By Sir Edmund G. Loder, Bart., Vice-President Z. S. 

[Received June 24, 1918 ; Read October 22, 1918.] 

Fro in the books on ISTatural History we have been given to 
understand that Beavers breed only once in the year, and that 
the young ones are born between the end of April and the 
beginning of June, after a period of gestation which is believed 
to last about fourteen weeks. 

In a book called ' In the Beaver World,' by E. A. Mills, it is 
stated that the number in a litter varies from one to eight, and 
that the eyes of the young ones are open from the beginning, 
and in less than two weeks they appear in the water accompanied 
by their mother. 

It is difficult to give an opinion as to how old the young- 
Beavers are when we first see them at Leonardslee. They are 
then about the size of rabbits, and we have supposed them to 
have been born six weeks or two months befoi-e, but we feel we 
have very little evidence to go on. I have always noticed the 
young ones of this size swimming about alone, the mother taking 
no notice of them. 

In January 1916 a pair of Canadian Beavers were received here. 
(I will call these No. 1 and No. 2.) 

On Dec. 11, 1916, a young beaver was seen, about the size of a 
rabbit. (I will call this one No. 4.) 

On July 10, 1917, three young ones were seen ; these again 
were about the same size as No. 4 when it was first seen. (I will 
call these three Nos. 6. 7, and 8.) 

Some time in August 1917 two young beavers were seen on 
the bank together. One was considerably larger than the other. 
The smaller one was recognized as No. 4, and we have to come to 
the conclusion that the other must have been born in the spring 
of 1916, soon after the arrival of the pair Nos. 1 and 2. (I will 
call this one No. 3.) 

On June 15, 1918, a young beaver was seen for the first time. 
Although only seen so lately, it is clear from its size that it 
must have been born some months ago, perhaps in December 
1917 or January 1918. (I call this one No. 5.) 

At the end of 1917 we had noticed that the old female seemed 
heavy in j^oung, and were rather disappointed not to have seen 
any signs of a litter, but it seems that she had one after all. 

On June 18, 1918, a very small beaver Avas seen (I call this 
one No. 9). It was not larger than a big rat. The little one 
was obviously out before the authorized time, for the mother (not 
the old female) went after it, and taking hold of a piece of its skin 
swam back with it to the mouth of the burrow, which is under 
water, and, letting go with her mouth, pushed it with her paws. 

I think this young mother must be No. 3, which I suppose was 



256 NOTES ON THE BEAVERS AT LEONARDSLEE. 

born in May or June 1916, therefore she was only just two years 
old when this little one was born. 

It will be interesting to see how long it will take the young 
one to grow to " rabbit " size. 

Now that there are two females bearing young ones, it will be 
difficult or impossible to make any exact observations. 

It is certain that the old female had young ones twice in one 
year: in October 1916 (seen December 1916) and in May 1917 
(seen July 1917), and it is probable that she had young ones on 
the following dates : — 

April 1916, 

October 1916, 
May 1917, 

December 1917. 

For some periods Beavers will show up continually in the day- 
light, and then will come an interval when they are seldom on 
view, or only one or two. 

When the pair of Canadian Beavers first arrived in January 
1916, I saw only one at a time for several months, until at last 
I was afraid that one must have died ; but, just as I had come to 
this conclusion, I saw the two together on the bank. 

To get accurate statistics is not so easy as might be imagined, 
but I think that the notes I have made are not far from the 
truth. 



MADAGASCAR FROGS OF THE GENUS MANTIDACTYLUS. 257 



16. On the Madagascar Frogs o£ the Genus Mantidactylus 
Blgr. By G. A. Boulenger, F.H.S., F.Z.S. 

[Received September 18, 1918 : Read October 22, 1918.] 
(Published by permission of the Trustees of the British Museum.) 

Index. 
Geographical : Page 

Madagascar Frogs 257 

Systematic : 

Mantidactylus, Synopsis of Species 258 

M. amhohimitomhi, sp. ii 260 

Agli/ptodactyliis, g. n., for Limnodytes niadagascariensis 

A. Dum 261 

Among the B:iany peculiar featui-es of the herpetological fauna 
of Madagascar is the fact of the genus Eana, so numerous in 
species in Continental Africa and the Indo-Malayan Region, 
liaving only two representatives : Ji. {Tonwpterna) lahrosa Cope, 
allied to the South African R. natcdensis A. Smith, and E. {Pty- 
chadena) mascareniensis D. & B., distributed over the greater 
part of Africa, the Seychelles and the Mascarenes included. 

Most of the Madagascar frogs originally referred to Bana or 
Liinnodytes {Hylorana) have proved to be distinguished by the 
presence of an intercalated bone between the penultimate and 
distal phalanges of the fingers and toes and have been referi-ed to 
an autochtonous genus, Mantidactylus *. In the species grouped 
by me under this genus, the swellings or discs in which the 
fingers and toes terminate bear on the lower surface a ring-shaped 
groove, defining a circular or transversely elliptic area, thus afford- 
ing a further distinctive character by which to recognise them 
among those species of Rana in which digital discs are likewise 
present. 

One species, Limnodytes viadagascariensis A. Dum. [R. ingioi- 
nalis Gthr.), which I had left in the genus Rana, has been shown 
by the late Dr. F. Mocquard to be also provided with the inter- 
calary phalanx and therefore i-eferred by him to Mantidactyhos ; 
but as in this species the small digital terminal expansions are 
devoid of the groove to which I now draw attention, I consider 
it to be entitled to generic distinction, under the new name of 
Aglyptodactylus. In this A. madagascariensis, the omosternum 
is forked at the ba§e, as in Mantidactylus, the nasal bones are 
small, oblique, and separated from each other as well as from the 
frontoparietals, and the terminal phalanges are obtuse ; there are 
no femoral glands. 

W6 are now acquainted with 22 species of Mantidactylus, to 
which a twenty-third is here added. A key to the identification 

* Ann. & Mag. N. H. (6) xv. 1895, p.- 450. 



258 MR. G. A. BOULENGER ON MADAGASCAR 

of the species was drawn up by Mocquard in 1909*, but as this 
key does not seem to me to work well and as the arrangement 
therein followed does not at all expi-ess the natural affinities, I 
have prepared a synopsis in which I have endeavoured to make 
good these deficiencies, so fai- as it is possible to do so in a linear 
sequence. 

Syno2)sis of the 6'/>ec/es o/"Mantidactylus. 

I. Glandular dorso-lateral fold, if present, not confluent with the supratemporal 
fold. 

A. Discs of fingers very small, usually smaller than those of the toes; snout 

rounded or very obtusely pointed, not or but feebly projecting beyond 
the mouth ; loreal region oblique ; belly perfectly smooth, or very feebly 
granulate behind. 

1. Toes entirely or nearly entirely webbed; head broader than long; back 

granulate. 
Tympanum hidden or small, very indistinct, and 

distant from the eye; tibio-tarsal articulation 

not reaching beyond the eye; heels meeting or 

not, when the limb is folded at right angles to 

the bodj' ; tibia 2^^ to 3 times as long as broad, 

2^ to 2| times in length from snout to vent M. guttitJatus Blgr. 1881 f. 

Tympanum distinct, small and distant from the 

eye; tibio-tarsal articulation reaching eye or tip 

of snout Jf. ^raMfZjf^i'eri Mocquard, 1895. 

Tympanum distinct, J diameter of eye ; tibio-tarsal 

articulation reaching posterior border of eye 31. inaudax Peracca, 1893. 

2. Toes i to § webbed ; head as long as broad ; tympanum very distinct, f to 

f diameter of eye. 

Series of vomerine teeth nearly equidistant from 

each other and from the ohoanse ; tibio-tarsal 

articulation reaching eye ; heels meeting ; tibia 

2f to 3 times as long as broad, 2\ to 2^ times 

in length from snout to vent ; back smooth M. ahitus Peracca, 1893. 

Series of vomerine teeth much nearer the choanoe 

than each other ; tibio-tarsal articulation reach- 
ing tip of snout, or between eye and tip of snout; 

heels strongly overlapping; tibia 3^ to 4 times 

as long as broad, If to 2 times in length from 

snout to vent ; back with glandular longitudinal 

folds M. betsileanus Blgr. 1882t. 

B. Discs of fingers small, as large as or larger than those of the toes; belly 

smooth or granulate only on the sides and behind. 

1. Head as long as broad or a little broader than long ; snout rounded or 
obtusely pointed; toes at least 3 webbed. 

a. Tibio-tarsal articulation reaching tympanum or posterior border of eye ; 
heels not overlapping; tibia 2\ to 3 times as long as broad, 2 to 
2f times in length from snout to vent ; first and second fingers equal; 
loreal region oblique ; back smooth or with indistinct flat warts ; belly 
smooth. 

Tympanum 2 to f diameter of ej'e ; toes f webbed. M. ciirtus Blgr. 1882. 
Tympanum f to f diameter of eye ; toes i to J 

webbed 31. biporus Blgr. 1889. 



* N. Arch. Mus. (5) i. p. 55. 

+ Includes 31. pic/er Mocquard, 1900. 

X Includes 3£. mtdtiplicatiis Boettg. 1913. 



FROGS Of THE GENUS MANTIDACTYLUS. 259 

I). Tibio-tarsal articulation reacliing eye or between eye and nostril ; toes 
f to f webbed ; tympanum f to once diameter of eye. 
tioreal region oblique ; tympanum not more than 
1^ times its distance from the eye ; tibia 2^ to 
3 times as long as broad ; inner metatarsal 
tubercle ij to j length of inner toe ; back with or 
without small elongate warts ; belly perfectly 

smooth .' Jf. amboJi imitombi, sp. n. 

Loreal region oblique ; tympanum 2 to 3 times its 
distance from the eye; tibia 3 to 8^ times as 
long as broad ; inner metatarsal tubercle ^ to j 
length of inner toe; back with elongate warts 
or glandular folds ; belly granulate on the sides 

aud behind M. ulcerosus Boettg. 1880. 

Loreal region nearly vertical ; inner metatarsal 
tuberele ^ length of inner toe; two glandular 

folds along the back M. hellyi Mocquard, 1895. 

c. Tibio-tarsal articulation reaching be- 
yond tip of snout; toes \ webbed; 
tympanum a little smaller than the 
eye; three glandular folds along the 

back ; belly smooth M. opiparis Peracca, 1893. 

3. Head a little longer than broad; snout pointed, stronglj' projecting beyond 
the mouth ; tibio-tarsal articulation reaching anterior border of eye or 
nostril; toes not more than^ webbed; belly smooth. 
Tympanum larger than the eye ; first finger 
shorter than second; two glandular folds along 

the back M. cerumnalls Peracca, 1893. 

Tympanum about i diameter of eye; first finger as 
long as or slightly shorter than second ; back [Hew. 1913. 

with large glands 31. glandulosus Meth. & 

C. Discs of fingers rather large, at least nearly twice as broad as the penultimate 
joint, as large as or larger than those of the toes. 

1. Belly perfectly smooth ; tibio-tarsal articulation reaching eye ; heels meet- 

ing ; tibia 3 times as long as broad, 2 to 2^ times in length from snout to 
vent ; toes entirely or nearly entirely webbed ; first finger much shorter 
than second ; loreal region nearly vertical. 
Head longer than broad ; snout pointed, strongly 
projecting beyond the mouth; tympanum £ 

diameter of eye If. wa/ori Blgr. 1896. 

Head as long as broad ; snout rounded or obtusely 
pointed, moderately projecting; tympanum \ to 
I diameter of eye M. cowanii Blgr. 1882. 

2. Belly granulate behind and on the sides onlj^ ; heels overlapping ; tibia 

3^ to 4 times as long as broad; toes f to nearly entirely webbed; inner 
metatarsal tubercle 5 to g^ length of inner toe ; first finger a little shorter 
than second ; t3anpanum | to f diameter of eye ; loreal region nearly 
vertical. 
Tibio-tarsal articulation reaching eye, nostril, or 

tip of snout; tibia If to 2 times in length from 

snout to Vent ; tongue usually with a conical 

papilla in the middle of the anterior third M. luguhris A. Dum. 1853*. 

Tibio-tarsal articulation reaching tip of snout or 

beyond ; tibia If times in length from snout to 

vent M.flavicrus Blgr. 1889. 

3. Belly granulate ; heels stronglj* overlapping. 

a. Tibio-tarsal articulation reaching eye or tip of snout ; tibia 3^ to 
4^ times as long as broad. If to 2 times in length from snout to vent ; 
tympanum i to f diameter of eye; loreal region nearly vertical. 
Toes f webbed ; inner metatarsal tubercle small, 
feebly prominent, \ length of inner toe ; a narrow 
dorso-lateral glandular fold M. granulatus Boettg. 1884. 

* Includes M. femoralis Blgr., 1882, and amhreensis Mocquard, 1895. 



260 MK. G. A. BOULENGER ON MADAGASCAR 

Toes i webbed ; inner metatarsal tubercle strong 

and prominent, compressed, 2 to | length of 

inner toe; no dorso-lateral fold M. redimitus B\gv. 1889. 

b. Tibio-tarsal articulation reaching beyond tip of snout; tibia 4^ to 
times as long as broad, Ij to If times in length from snout to vent ; 
loreal region oblique. 
Toes nearly entirely webbed ; inner metatarsal 

tubercle ^ length of inner toe ; tympanum i to 

t diameter of eye ; a pair of inwardly curved 

glandular folds on the anterior third of the back, 

from the upper eyelids ; heel with a dermal pro- 
cess or spur ^^- Jiiteus Metli. & Hew. 1913. 

Toes i to f webbed ; inner metatarsal tubercle k to 

I length of inner toe ; tympanum i to | diameter 

of ej-e ; a curved glandular fold on eacli side 

from the upper eyelid to between the shoulders, 

followed by a straight fold -M". pUciferus Blgr. 1882. 

Toes \ webbed ; inner metatarsal tubercle f to J 

length of inner toe ; tympanum f to J diameter 

of eye ; upper parts rough with prominent glan- 
dular folds and tubercles; heel with a dermal 

process or spur Jf. orspej- Blgr. 1882. 

II. Glandular dorso-lateral fold extending from 
behind the eye to the hip ; loreal region ver- 
tical ; tympanum f to once diameter of eye ; 
discs of fingers and toes rather large ; tibio- 
tarsal articulation reaching tip of snout or a 
little beyond ; tibia 4| to 5 times as long as 
broad, If to 1^ times in length from snout to 
vent; toes i webbed ; belly granulate behind. M. albofrenatns F. Miill. 1892*. 

Nothing is known of tlie development and larvte of these frogs, 
but the eggs are remarkably large, measuring 5 mm. in diameter 
in M. guttidatus ($ 120 mm. from snout to vent), 3 mm. in 
M. luguhris ( $ 50 mm.), 2-5 mm. in M. betsileamcs ( $ 33 mm.). 

Mantidactylus ambohimitombi, sp. n. 

Vomerine teeth in short transverse or oblique series behind the 
level of the choanal, equidistant from the latter and from each 
other. Head a little broader than long ; snout rounded, feebly 
proiecting beyond the mouth, with indistinct can thus and very 
oblique, concave loreal region ; nostril equidistant from the eye 
and from the end of the snout ; interorbital region as broad as or 
a little narrower than the upper eyelid ; tympanum distinct, f to 
f the diameter of the eye, 1 to IJ times its distance from the 
latter. Fingers moderately long, first and second equal or first a 
little the longer, the discs small, not very much larger than the 
well-developed subarticular tubercles. Toes moderately long, 
f webbed, the discs about as large as those of the fingers ; no 
tarsal fold ; inner metatarsal tubercle oval, moderately promi- 
nent, i to I the length of the inner toe; no outer tubercle. 
Tibio-tarsal a,rticulation reaching the eye or betAveen the eye and 
the nostril ; tibia 2| to 3 times as long as broad, 14- to 2 times in 

* M.frenatus Boettg., 1913, is probably identical with this species, although the 
hind limb is longer and the discs of the fingers and toes are described as very small. 



FROGS OF THE GENUS MANTIDACTYLUS. 261 

length from snout to vent, as long as or a little shorter than the 
foot. Skin finely granulate above, with or without elongate flat 
warts on the sides of the body ; a strong, curved glandular fold 
from the eye to the shoulder ; lower pai'ts smooth ; femoral gland 
more or less distinct, with a single pit, or absent. Brown above, 
spotted or mai'bled with daikei', often with a large dark trian- 
gular spot between the eyes ; a dark canthal stieak and a tem- 
poral band, usually light-edged beneath ; a yellow vertebral streak 
sometimes present ; limbs with more or less distinct dark cross- 
bands ; hinder side of thighs usually dark brown, with small 
yellow spots. White beneath, unifoi'm or mottled with brown, 
or nearly entirely brown. Male without vocal sacs. 

Nasal bones rather large, narrowly separated from each other 
and from the frontoparietals. 

From snout to vent 65 millim. 

Several specimens from the Ambohimitombo Forest, Mada- 
gascar, from the collection of Dr. Forsyth Major, 1896. 



Proc. Zool. Soc.~1919, No. XIX. 19 



CILIARY ACTION IN PLEUROBRACHIA PILEUS. 263 

17 Ciliary Action in the Internal Cavities oE the C'tenophore 
Pleurohrachia pileiis Fabr. Bv James F. Gemmill, 
M.A., M.D., D.Sc, F.Z.S. 

[Received October 2, 1918: Read November 5, 1918.] 
(Text-figures 1 & 2.) 

During life ciliation is active throughout the internal cavities 
of Pleurohrachia ; and the latter are wide enough to allow the 
dii-ection of the ciliary action on their different surfaces to be 
made out from the motion of particles suspended in the con- 
tained fluid, e. g. small oil globules, alimentary particles, and 
debris. 

The "circulation" * is an extremely orderly one and meets the 
physiological need for continuous regulated change through the 
whole of the internal cavities. 

, On the whole the circulation inside the funnel system goes 
on independently of that within the stomodseum t- So far as I 
could make out, except under the influence of peristaltic action, 
only slight interchange of fluid between the stomodfeum and the 
funnel takes place. 

T. StomodcBimi (text-figs. 1 & 2), 

Round the margin of the mouth there is a very narrow band 
best seen in young specimens, the cilia of which stiike into 
the mouth-cavity. Up the middle of each lateral wall of the 
stomodfeum, and continued on the thickenings in this region, and 
to the infundibular opening, there is a track with aboralward 
ciliation. Over the rest of the lateral wall of the stomodseum the 
ciUation is oralwards with a slant towards the sagittal angles. 
' Along the sagittal angles from the opening of the funnel to that 
of the mouth the ciliation is stronger and in the oralward 
direction. 

II, Funnel and Canal System (text-fig. 2). 

We may best follow the circulation here by beginning in the 
floor (oral wall) of the funnel at points on opposits sides of the 
opening from stomodsBum into funnel. It will be remembered 
that the aboralward currents up the middle of the sides of the 
stomodaeum lead to these points. Working transversely outwards 

* Reference may be made to the following recent papers on ciliation : — 

Carlgren, 0. Biol. Centralbl. xxv. 1905, pp. 308-322 (Actinians, Madreporarians). 
Oi-ton, J. H. Journ. Mar. Biol. Assoc. U. K. ix. 1912, pp. 144-478 (Ascidians, 

Molluscs). 
Orton, J. H. Ibid. x. 1913, pp. 19-49 (AmpJiioxus, Ascidians, Molluscs), 
(iemmill, J. F. Proc. Zool. Soc. Lond. 1915, pp. 1-19 (Starfisb). 
Widmark, E. M. P. Zs. Allg. Phys. Jena, xv. 1913, pp. 33-48 {Aurelia aurita). 
t In this paper the whole of the cavity between mouth and funnel-opening is 
called stomodseura. 

19* 



264 



DR. J. F. GKMMILL ON CILIARY ACTION IN THE 



Text-fiafure 1. 




Diagram of lateral wall of stomodseum of Fleurohraehia, showing direction of 
ciliary currents. 

If . 0., mouth-opening ; i^.O., funnel-opening. 

(For explanation see text.) 

Text -figure 2. 




Diagram of internal cavities of Fleurohracliia (transverse or infundibular plane in 
plane of paper). The arrow-heads in the "walls of the cavities point in the 
direction of the ciliarj^ currents. 

1. StomodsQum. S- Funnel. 3. Paragastric canal. 4- Tentacular canal. 
5. Perradial canal. G. luterradial canal. 7. A sub-sagittal canal. 8. A sub- 
transverse canal. 9. Adradial canal. 



INTERNAL CAVITIES OF PLEUROBRACIIIA PILEUS. 265 

from either of the points in question we find the current strongly 
outwards towards the equator, in the floor of the funnel, but soon 
meet the opening into the paragastric canal. The axial wall of 
this canal carries cilia which strike oralwaxds, while on its abaxial 
wall the cilifi strike in the aboral direction. The canal is thus 
bathed mesially by an in-going and laterally by an out-going cur- 
rent (text-fig. 2, 3). Exactly the same thing holds good for the 
ciliation and currents within the tentacle-canals (text-fig. 2, 4)- 
On the floor (oral wall) of the perradial, interradial, and adradial 
canals the ciliation is outwards, i. e., towards the entrances into 
the meridional canals. 

The adjacent halves of neighbouring sub-sagittal canals are 
ciliated along the whole length of their axial walls in the oral- 
ward direction. This holds good also for the adjacent halves 
of neighbouring sub-transverse canals. The areas of oralward 
ciliation thus correspond with the distribution described for the 
female gonads at the sides of the canals. The axial walls of the 
remaining halves of all the meridional canals {cf. distribution of 
tihe male gonads) are ciliated in the aboralward direction. 

Along the roof (aboral wall) of the adradial, interradial, and 
perradial canals the ciliation is inwards, i. e., towards the main 
cavity of the funnel. The walls of the aboral exten-sion of the 
funnel have an aboralward ciliation, while an oralward reflux 
takes place down the middle of this extension into the main 
cavity of the funnel. 

Here mixing occurs and currents are caught up anew into the 
paragastric, tentacle, and other canals. 

The arrangement above described proved constant for a con- 
siderable number of large and small PleitrohracMa examined. 
The tissues ai'e transparent enough to allow the examination to 
be made in undissected specimens with the help of a binocular 
microscope. Fine carmine grains can be used to supplement the 
evidence of the particles floating in the gastrovascular fluids. 
Mixed with sea-water the powdered carmine will occasionally be 
ingested into the stomodaann by natural peristaltic action, or it 
may be injected into this cavity with the help of a pipette. 
After a short interval the carmine is expelled from the mouth, 
but meantime in successful cases sufficient particles to be visible 
have found their way into the funnel and the cavities leading 
therefrom. 

The specimens were obtained last June at the Millport 
Biological Station, and the work was done partly there and 
partly at Glasgow University, the cost of obtaining material 
being met out of a grant from the Carnegie Scottish Universities 
Trust. 



CAPT. D. M. S. AVATSON ON SEYMOURIA. 267 

18. On Set/ inouri a, the most primitive known reptile. By 
D. M. S. Watson, M.Sc, Capt. U.A.F,, Lecturer in 
Vertebrate Palseontologj, University College, London. 

[Received September 9, 1918 : Read Novembev 19, 1918.] 
(Text-figures 1-15.) 

The reptile Seijmouria hayloriensis was originally described 
from two imperfect skulls, one in connection with a few vertebrae 
and the shoulder-girdle, by Prof. Broili of Munich, whose excel- 
lent description made us well acquainted with the structure of 
the upper and lateral surfaces of the greater part of the skull, and 
gave general information about the palate, occiput, and anterior 
axial skeleton. 

In 1910 Williston described as Besmospondylus anomalus a 
collection of vertebrae, a humerus, femur, and some other bones, 
which he subsequently recognised as belonging to a young Sey- 
moitria. In 1911 the same author described a beautifully 
complete skeleton, publishing a restoration to whose accuracy I 
am glad to be able to bear testimony. At the same time he 
suggested that the skull described by Cope as Conodectes favosus 
was really Seymouria, a determination which is undoubtedly 
correct. As Cope's description of Conodectes is quite insufficient 
for recognition of the skull, and Seymoitria is a name universally 
known, I propose to regard Conodectes as a nomen oiudtim and 
relegate it to the synonymy. 

In 1914, V. Huene published figures of the type skull of 
Conodectes, but added nothing to our knowledge; and in 1915 
I gave a short description of such knowledge of the otic region 
as covild be obtained from the rather badly preserved skull of 
Conodectes. 

In danuai-y 1914, through the kindness of Prof. Broili, I was 
able to make a careful examination and drawings of the type 
material of Seymoiiria in Munich, which at that time, owing to 
a new and more complete preparation, showed many features of 
the structure of the palate and occiput which were not referred 
to in the original description. 

In 1915 I was so fortunate as to collect some Seymouria 
material in Texas. The most important of my specimens was 
found weathered out on the side of a small hillock about 20 yards 
away from the Cradock bone-bed quarry. I at first supposed it 
to be a single individual, but have subsequently found that two 
are represented. The better individual is represented by the 
pelvis, both hind legs, fifteen presacral, the sacral and the caudal 
vertebrae in a connected series, and many ribs ; these bones are 
connected by matrix and are all articulated. Almost certainly 
belonging to this individual are the atlas and axis and three 



268 CAPT. D. M. S. WATSON ON SEYMOUKIA, 

succeeding vertebra3, with the incomplete right scapula, coracoid, 
and clavicle attached by matrix, the occipital region and part of 
the right side of the skull and lower jaw, the lower end of a 
radius and ulna, and a metacarpus. The associated skeleton is 
represented by a saci^al and nine presacral vertebrae, an incom- 
plete femur and ischium. 

Another specimen collected on West Coffee Creek consists of 
numerous fragmentary bones of a young individual washed 
perfectly clean. I also obtained certain isolated bones. 

This material and the new preparation of that at Munich 
allow me to add materially to our knowledge of the structure of 
this most interesting form, perhaps the most perfect annectant 
type known to us. 

Skull. 

Profs. Broili and Williston have given a satisfactory account 
of the upper and lateral surfaces of the skull. The Munich 
skulls and that of my skeleton give an equally complete know- 
ledge of the occiput and palate. For reference I shall refer to 
that Munich skull which has a shoulder-girdle belonging to it 
as A, the other as B. 

Basioccijntal. — This bone is very well shown in my skull. It 
is a small bone, remarkably thin dorso-ventrally and of consider- 
able width. The doi'sal surface appears to be completely covered 
by the exoccipitals and forms no part of the floor of the brain- 
cavity. The posterior surface forms part of the condyle, which 
is wide from side to side, shallow and rounded. The ventral 
surface is widely exposed as a quadrangular area bounded in 
front by the suture with the basisphenoid. The posterior part 
of the lateral border is in contact with the exoccipitals, in advance 
of which it is cut out into a shallow notch, the lower border of 
the fenestra ovalis. Anteriorly the lateral margin is produced 
downward and outward to take part in the formation of the 
tuber basisphenoidalis. 

Basisphenoid. — The basisphenoid is completely known, so far 
as concerns its inferior surface, from the Munich skulls and that 
of my skeleton. 

Posteriorly it has a suture with the basioccipital, the ends of 
which lie on the summits of the well-marked tubera. From the 
tubera a pair of prominent ridges run forward on the ventral 
surface, so that this face forms a smooth concavity between them. 
At the level of the basipterygoid processes these ridges die out, 
so that the surface becomes gently convex. The basipterygoid 
processes (shown in Munich A & B) are extremely short and end 
in a flat articular surface, w^hich does not directly support the 
pterygoid but is attached to a separate small bone very clearly 
shown in Munich B. The lateral surface of the basisphenoid, 
between the tuber and the basipterygoid pi'ocess, is concave and 
passes indistinguishably into that of the prootic. About three 
millimetres above the lower edge of the tubera a minute foramen 



THE MOST PRIMITIVE KNOWN REPTILE. 



269 



opens out from the basisphenoid. A broken face about 5 mm. in 
advance of the tubera shows that in that region the basisphenoid 
is a thin plate scarcely a millimetre thick medially, 

Text-fieure 1. 




Seymouria hayloriensis Broili. — Restoration of the palate, X f . 

General shape and palate from the types in Munich. Details of maxillary dentition 

from fragments in my possession. Basisphenoid and occipital region from 

mj' skeleton. 
Reference letters -.—B.Oc, basioccipital ; B.Sp., basisphenoid; Ect.Pt., ectoptery- 

goid; Ex.Oc.,exoccipital; Mx., maxilla ; P.Mx., premaxilla; Pal., palatine ; 

Par.Oc, paroccipital ; Pr.O., prootic ; Pk.V., prevomer; Qu., quadrate; 

X, autogenous basipterygoid process ? 

Parasphenoid. — The parasphenoid is well shown_ in Munich B 
as a short narrow rostrum projecting forward in the palate 
between the pterygoids. 

The type of Conodectes shows that it supports a large ethmoidal 
ossification which surrounds the anterior part of the brain. 

Exoccipitcd. — The right exoccipital is well shown in my skull, 



270 



CAPT. D. M. S. WATSON ON SEYMOURIA, 



in which unfortunately a, piece about 2*5 mm. thick is missing 
from the middle of its height, this not having been collected. 

The exoccipital rests on the upper and lateral surface of the 
basioccipital, meeting its fellow above that bone so as to form 
the floor of the brain-cavity in the occipital region. Posteriorly 
it projects behind the basioccipital and is provided with an arti- 
cular face looking downwards and backwards which forms a.bout 
a quadrant of the occipital condyle. Above the body the bone is 
continued up so as to form the side wall of the brain-cavity until 
its posterior surface is ovei'lapped by the occipital flange of the 
dermo-supraoccipital. 

On the inner surface the exoccipital is excavated into a deep 
pit from which a rather laige hypoglossal foramen starts to pass 
through the bone and open into the large vagus foramen. In 

Text-figure 2. 




Qv. Pr. ParOc Ex.Oc.B.0c.Fen.0v. QuJ, 



Seymouria hayloriensis Broili. — Restoration of the occipital view, X f , 

Membraiie-boues from Munich A. Quadrate from a young individual collected 
by the writer. Occiput from my skeleton. 

Reference letters as before with i^D.S.Oc, dermo-supraoccipital ; Fen.Ov., fenestra 
ovalis; Qu.J., quadrato-jugal; Sq., squamosal ; Tab., tabular. 



fi'ont of the ridge which bounds the hypoglossal foramen 
anteriorly, the inner surface of the exoccipital turns outward 
and forms the posterior margin of the foiamen for the Xth 
nerve. Tlie upper end of the exoccipital is fused with the par- 
occipital, no suture being visible. It approaches its fellow of the 
opposite side over the foramen magnum, but leaves a space for 
the cartilaginous supraoccipital. In my skull the space where 
the supraoccipital should be is entirely free from bone, but in the 
type of Conodectes there is some evidence of a slight supra- 
occipital ossification. 

Paroccipital. — The paroccipital is fused with the exoccipital on 
the bacls; of the skull. It is separated from the lower part of 
that bone by a large notch, the foramen for the vagus nerve. 



THE MOST PRIMITIVE KNOWN REPTILE. 



271 



Laterally to this it forms a deep plate extending downwards 
nearly to the level of the ventral surface of the basioccipital, 
where it ends in a suture with the prootic laterally and in a free 
margin, part of the border of the fenestra ovalis, at its inner end. 
The upper outer part of the posterior surface is overlapped by the 
occipital flange of the tabular. Between this region and the top 
of the exoccipital the upper pai-t of the posterior surface is turned 
forward so as to form a groove, the lower part of the post-temporal 
fossa. The cranial end of the paroccipital appears to be narrow, 
so that the inner ear is widely open to the cranial cavity in the 
bony skull. 

Prootic. — The prootic is fused with the lateral margin of the 
basisphenoid below and has a suture with the paroccipital 
behind. Its lateral face, which alone is satisfactorily shown, has 

Text-figure 3. 



Q5,0c, TftB 




Ou, Pt f:b.Pt 



Seymouria hayloriensis Broili. — Occipital view of the type-skull 
Munich A, X f . 

Refeceiice letters as before with : — F.B.Pe., facet on the pterygoid for articulation 
with the "basipterygoid process." 



a powerful ridge running horizontally along it at about the 
middle of its height. Below this it is concave, its lower border 
forming part of the rim of the fenestra ovalis mesially and laterally 
uniting with the paroccipital, the joint bones forming the massive 
paroccipital process which in section has a V-shaped lower surface. 
Above the ridge the prootic is essentially flat, having however 
a deep groove leading to a foramen towards the outer end. The 
anterior edge of the bone is damaged, but the minute foramen fqr 
the facial nerve is just preserved. 

The endocranial end of the bone is shown to be formed by an 
extremely thin shell. 

Dermo-sujyraoccijyital and Tabular. — The dorsal exposure of 
these bones on the upper table of the skull has been accurately 
described by Broili. Munich A shows their occipital aspect 
perfectly. Each dermo-supraoccipital has a small descending 



272 



OAPT, D. M. S. WATSON ON SEYMOURIA, 



lappet passing vertically flownward, the inner border of this 
forms part of the margin of the foramen magnum, the outer, that 
of the post- temporal fossa. 

The tabular has a similar occipital flange forming the outer 
margin of the post-temporal fossa and covering a large area of 
the posterior surface of the paroccipital. 

My skull suggests the presence of some sort of a flange on the 
ventral surface of the tabular articulating with the end of the 
paroccipital process. 

Text-figure 4. 




Seyniouria hayloriensis Broili. — Palate, X f. Drawn from Munich A. 

Reference letters as before with : — B.Pt., the basipterygoid articulations on the 
basisplienoid ; R.Pal.Th., right palatine tusk. 



Pterygoid. — The pterygoid is completely exposed in Munich A. 

The quadrate ramus forms a verticall3? standing plate whose 
posterior surface is concave. The outer margin has a long- 
suture with the squamosal below tlie otic notch, veutrally it 
separates from that bone so as to leave exposed an area of the 
quadrate, with which bone it has a large powerful suture. 

The inner margin of the quadrate ramus approaches the 
anterior border of the prootic closely, the bone in this region 
though deep not nearly reaching the skull-roof. This part of the 
ramus bears towards its ventral edge a large, well-developed, 
slightly cupped articular facet. 

When the perfectly preserved pterygoids and basisphenoids of 
Munich A and B are compared, it is seen that there is a large 



THE MOST PRIMITIVE KNOWN REPTILE. 



273 



triangular space between this facet on the ptei'ygoid and that on 
the basipterygoid process with which in ordinary reptiles it 
should articulate. In Munich B, this space is filled up by a 
small separate bone on the right side of the skull. The fact that 
although in A this bone is not preserved, the facets on the 
pterygoids and basi pterygoids are identical in the two specimens, 
shows beyond dispute that it is a separate bone. Nothing similar 
is known in any other Tetrapod. 

Text-fiffure 5. 




Seymouria hayloriensis Broili. — Palate, X |. Drawn from Munich B. 
Reference letters as before with : — Pae.Sp., pavasphenoid. 



In advance oi the quadrate ramus the pterygoid is represented 
by a lai'ge flat bone in the palate which articulates with its 
fellow in the middle line in advance of the parasphenoid, and 
laterally has sutures with the transverse, palatine, and prevomer. 
That margin of the palatal part of the pterygoid which forms the 
anterior limit of the subtemporal fossa is slightly deflected 
laterally, so that with the ectopterygoid it forms a small flange 
against the inner surface of the lower jaw. 

Ectopterygoidj. — The ectopterygoid is a bone of few features. 
It forms merely a plate in the palate articulating with the 
pterygoid in a suture only shown incompletely in my skull, a 
long suture with the palatine and presumably another with the 
maxilla. About the middle of its area in Munich A is a small 
depression probably intended to receive a tooth on the lower 

Palatine. — The palatine is well shown in Munich A and in my 



274 CAPT. D, M. S. WATSON ON SEYMOURIA, 

skull. It has a long suture with the pterygoid, is bounded 
posteriorly by the transverse, and in front forms certainly the 
back and appai-ently also a large part of the lateral margin of 
the internal narial opening. 

Its most interesting feature is the presence of a pit surrounded 
by an upstanding ridge, lying on the palatal surface just posterior 
to the nostril. This pit contains two lai-ge tusks, which replace 
one another so that normally only one is functional at once. 
The whole arrangement is identical with the large palatine tusks 
and their pits iru Labyrinthodonts. It is remarkable that this 
tooth has only been described by Cope in Conodectes. It is well 
shown also in Munich A and my skull. 

Prevomer. — Only the posterior end of the prevomer is known. 
The two separate the internal nares and articulate with the 
anterior ends of the pterygoids. 

Qtmdrate. — An incomplete but isolated and extremely well- 
preserved left quadrate belongs to my young West Coffee Creek 
specimen-. It has a well-ossified and rounded articular margin 
rather conspicuously divided into two condyles. The anterior 
face is concave. The outer side is entirely occupied by a sutural 
surface for the quadrato-jugal and squamosal, and the inner 
retains the impression of the tip of the quadrate ramus of the 
pterygoid. The exposed posterior surface is triangular and bears 
an irregular knob. 

Ductus naso-lachrymalis. — My skull shows quite clearly the 
presence of a naso-lachrymal duct lying within the substance of 
the lachrymal and extending from the orbit to the nostril. 

Septomaxilla. — Whilst removing the matrix from the sym- 
physial region of the mandible of my skull, I found a single 
septomaxilla which had obviously dropped down through the 
posterior naris. 

This element is a thin plate of bone bent round so as to clasp 
Jacobson's organ, and has a flat face for articulation with the 
dorsal surface of the prevomer. 

Lower jaw. — My skeleton retains the right ramus of the lower 
jaw from the symphysis to behind the anterior end of the supra- 
meckelian fossa. Both inner and outer sui'faces are well exposed, 
and a fortunate fracture along a horizontal plane lying just above 
the upper surface of the cavity of the jaw, which passes through 
the dentary and the three coronoids, places the structure beyond 
doubt, as it permits a definite distinction between sutures and 
cracks. The sutures are usually very visible, being filled with red 
iron oxide and the bone white. 

Articidar. — Both articulars of the yoimg specimen from Coffee 
Creek are preserved. 

The bone is very short and ends anteriorly in a flat face con- 
tinued in life by the remains of Meckel's cartilage. 

The articvilar surface is convex from back to front and is 
divided into two areas, corresponding to the condyles of the 
quadrate, by a low ridge running obliquely across. 

The outer face has a deeply depressed sutural area for the 



THE iMOST PRIMITIVE- KNOWN REPTILE. 

Text-figure 6. 



275 



D. 

Cor I^^-'>Den. 






5. 



Den. 

"''''" CorI 
5up,flN(;^ 



CoR.I. PalTh CorII. CorI 





5r 



PoST.Sp Pr.Rrt. f\N^. ^' 



Seymouria hayloriensis Broili.— Eight ramus of the lower jaw of my 
skeleton, X 1. 

A. Inner surface. The area covered with parallel oblique lines is covered with 

matrix; the teeth lu the dentary are restored on the evidence of fragments of 
that bone of a youug individual. The articular is drawn from that of a 
young individual and is probably slightly too small. 

B. Ventral view of a horizontal fracture passing just above the cavity of the lower 

jaw of my skeleton. 

C. Vertical section seen on the broken posterior end of the jaw. 

D. Vertical section seen on a fracture through the splenial region. 

Reference letters:— Ang., angular; Aet., articular; CoK. 1, II, III, the 1st 2nd 
and 3rd coronoids; Den., dentary ; Post.Sp., post-splenial ; Pe.Aet' pre- 
articular; Pai.Th., the palatine tusks, represented in dotted lines to 'show 
theiv position with the mouth closed ; Sp., splenial ; Sur.Ang., surangular 



276 . CAPT. D. M. S. WATSON ON SEYMOURIA, 

surangular ; the inner, more faintly marked depressions for the 
prearticiilar and angular. The posterior surface bears a minute 
knob representing the postarticular process. 

Dentary. — In general the sutures on the outer surface of the 
jaw are not well shown. The dentary, however, forms the entire 
outer surface for some distance behind the symphysis, posteriorly 
it ends in a point which is received in a groove in the surangular. 
Fragments belonging to the young individual show that it caj-'ried 
a single series of rather large round teeth. Its admesial surface 
has a long perfectly straight suture with the coronoids. 

Splenial. — The splenial appears to enter the symphysis. It 
forms the lower border of the anterior portion of the jaw, its 
suture with the dentary running along just above the- lower edge 
on the outer surface. The greater part of the bone, however, 
forms a flat inner surface articulating above with the first and 
second coronoids, and behind with the prearticular and pre- 
angiilar. 

Preangidar. — The preangular is a channel-shaped bone forming 
the lower part of the jaw, articvilating above with the dentary 
on the outer surface and the prearticular on the inner, with the 
splenial in front and the angular behind. 

Angular. — The angular is only very incompletely preserved, 
but is a bone of the ordinary reptilian pattern. 

Prearticular. — The prearticular is a very large bone running 
from the articvilar to the splenial. Its lower edge has long 
sutures with the angular and preangular, interrupted so as to 
leave an anterior internal mandibular vacuity between the pre- 
articular and pi-eangular and a posterior vacuity bounded by 
prearticular and angular. 

Coronoids. — The fracture referred to above shows that there 
are three coronoids forming together a continuous strip wedged 
in between the dentary and surangular on the outside and the 
prearticular and splenial within. 

The 1st or anterior bone is small and incompletely exposed, it 
is not shown to bear any teeth . 

The 2nd is of considerable size, and in the middle of its length, 
which in life lies immediately outside the palatine tusks, is 
thickened and carries a 'tightly packed mass of small granular 
teeth . 

The 3rd, posterior, coronoid forms the anterior border of the 
supra-meckelian vacuity, extending back along the sides of that 
opening in contact with the inner surface of the surangular 
outside and the upper edge of the prearticular below. This 3rd 
coronoid bears one large bluntly pointed tusk at about the middle 
of its length : this tooth is oval in cross-section and was appa- 
rently received in a special pit in the ectopterygoid. 

Vertebral Column. 

Atlas. — The first vertebra preserved in my skeleton is the 
atlas. The parts preserved are slightly displaced. 



THE MOST PRIMITIVE KNOWN REPTILE. 



277 



There is a large hemicylindrical intercentrum whose width 
agrees with that of the basioccipital part of the condyle. The 
lower surface is smooth, short blunt-ended backwardly directed 
processes forming its latei-al margin. The upper surface of this 
bone has laterally two large flat triangular areas for articulation 
with the neural elements; between these areas the anterior end 
of the bone is excavated into a conical pit, only half of which is 
present, but which resembles exactly that in the end of an 
ordinary centrum. 

Text-figure 7. 




IN 



C OakC^ C^ WC^ C 



8eymouria-hayloriensis Broili.— Three anterior vertebrae, X 2. 
Right lateral aspect. Left surface. 

The atlas is composed of drawings of the elements in my skeleton replaced in what 
appears to be their natural relations, the centrum of the axis and the inter- 
centrum before it are hypothetical. 

C2, &, centra of 2nd & 3rd vertebrae; In.C.i>2, 3^ intercentra 1, 2, & 3; 
Od., odontoid ; N' & N^, neural arches of 1st & 2nd vertebrae. 



One pair of other elements is preserved. Each of these has 
a high laterally compressed but antero-posteriorly elongated 
neural spine, whose posterior margin descends directly to the 
hinder end of a well-formed posterior zygapophysis. The 
anterior margin of the spine descends vertically about to the 
level of this articular face, and then turns forward so as to 
lie horizontally and form the upper edge of a squarish area of 
bone whose front edge is rounded and articular and of consider- 
able width. The lower part of this region ends in a short blunt 
point. It is obvious that the neural spines of the pair of elements 
lay in contact with one another and that the zygapophyses were 
supported by those of the succeeding vertebra. The anterior end 
could have articulated with the exoccipital part of the condyle, 
the intercentrum supporting the basioccipital part. 

Proc. Zool. See— 1918, No. XX. 20 



278 



CAPT. D. M. S. WATSON ON SEYMOURIA, 



The odontokl is a single bone. Its upper surface is slightly 
rounded but presents no trace of any articulation. The posterior 
surface is not exposed. The anterior surface is slightly convex, 
rising to a small point. This surface is in general triangular 
with the point at the mid-ventral surface. The lateral edges are 
slightly emarginate at about the middle of their height so as to 
divide the whole area into three ; of these, the lowest articulates 
with the anterior intercentrum, and the two dorsal with the 
neural arches. The lateral surface is nearly fiat, but rises some- 
wha.t towards the ends ; it is triangular, so that there must be a 
gap for a triangular intercentrum between the odontoid and the 
axial centrum. 



Text-figure 8. 



f\. 




D. 




D. 




F i--.-M!l:\! 




Setjmonria haijloriensis Broili. X 2.- 

A. Odontoid from in front-. 

B. Atlantal intercentrum from in front. 

C. Atlantal intercentrum from below, anterior end upward. 

D. Third intercentrum and centrum from below. 

E. Fourth centrum and fifth intercentrum. 



Axis. — The second vertebra preserved difiers in no important 
respect from the third and fourth. It has large well-formed and 
somewhat swollen zygapophyses, which are not very much pro- 
duced laterally, the lateral part of the neuiul arch immediately 
below the articular surface beini>- extended outward into a shoit 



THE MOST PRIMITIVE KNOWN REPTILE. 279 

but well-formed transverse process bearing a rib-facet distally. 
The neural spine of this vertebra is destroyed. 

The restoration (text-fig. 7) is only uncertain in the shape 
and size of the intercentrum between the odontoid and axial 
centrum and of that element itself. The sutures between the 
neural arch and centrum of the axis are obliterated. 

The 3rd and 4th vertebrae exactly i^esemble the 2nd, except that 
the arches are wider and the transverse processes longer. The 
neural spine is represented only by a little ridge. 

The 5th vertebra exactly resembles that in front of it except 
that it has a low but very massive neural spine. 

The centra of the 3rd, 4th, and 5th vertebrae are short antero- 
posteriorly and have a very well-marked keel, their sides being- 
excavated into deep concavities on each side of the middle line. 
The intercentrum between the 4th and 5th vertebrae is well 
displayed, it is nearly as long as the centrum before it, and 
laterally is carried out into well-marked processes for articulation 
with the capitula of the ribs. 

The next vertebra, completely preserved is the 9th, from which 
the series is complete and naturally articulated to the 10th 
caudal. 

The 9th is a typical Seymouina vertebra with a small well- 
rounded centri\m, considerably longer than that of the 5th. The 
arch is enormously wide and massive, but the ends of the com- 
paratively slender transverse processes project well bej'^ond the 
extremities of the zygapophyses. The 9th vertebra has a low but 
very massive spine whose summit is bifid. 

From this vertebra to the sacrum the sti'ucture of the vertebrae 
remains very unifoi-m, the only changes being that the transverse 
process gradually shortens, the spine becomes lower and more 
slender, and the ventral surface of the centrum becomes flattened. 

The 10th vertebra is unique in that it is entirely devoid of the 
slightest trace of a spine. I myself removed the matrix which 
covered it, and the surface is still practically perfect. 

My skeleton is abnormal in that the 23rd vertebra is asym- 
metrical, on the left side agreeing exactly with that in front, 
whilst the right side of the neural arch is much elongated and 
carries a sacral rib. 

The 24th vertebra, the sacral, is only exposed on the left side, 
•where it carries a veiy massive sacral rib, applied to the inner 
aspect of the ilium in the usual way. This vertebra has a rela- 
tively high and massive spine. 

As Prof. Williston has already remarked, the first caudal difliers 
markedly from all that precede it in its narrow and unexpanded 
neural arch. It has a fairly long transverse process. From hei'e 
to the sixth the caudals do not difier much, except that the spine 
gets progressively more and more slender and leans more steeply 
backward . 

The seventh and succeeding caudals bear no ribs and the 
transverse process has disappeared from them ; their centra are 

20* 



280 



CAPT. D. M. S. WATSON ON SEYMOURIA, 



small and liourglass-sliaped and separated from one another by 
an interval of more than their own length. The neural arch 
rests on the anterior part of the centrum and presents a large 
articular face to the space between the centra ; the lower part of 
this space m in part filled, up by the top of the chevron, but there 
can be no doubt that the condition in life was essentially embolo- 
merous, the cartilaginous iiitercentrum not having ossified. 

Text-figure 9. 




w 



IM 



CavN 



QwY 



Seymouria hayloriensis Broili. Ribs, X 1. 

The numbers are those which the bones drawn occupy in the series. 
Cau. IV. & V. are caudal ribs. 



72i&s.— The general distribution and structure of the ribs have 
been well described by Williston, to whose account, however, I 
am able to add some details of interest. 

The long slender atlantal rib is double-headed, the capitulum 
and tuberculum being carried at the ends of two widely separated 
branches. 

The upper part of the 5th rib, which has a very expanded 
distal end, is extraordinarily deep and massive. The two heads 
are widely separated, the (.-apitulum articulating with a special 
process on the intercentrum. 

From here backwards the ribs are more slender, and in the 
middle of the back the tuberculum is merely a facet on the upper 
edge of the rib, and not carried out as a special process. 

The sacral ribs cannot be described in detail. The caudal ribs 
are remarkably long and massive, they are well curved, and if 
found separately \\ould have been regarded as doi-sals. 



'rUB MOST PRIMITIVE KNOWN REPTILE. 



281 



Tlie ribs on the 4th and 5th caudals ar.e very distinctly double- 
headed, having a distinct emargination between the well-marked 
tuberculum and capitulum. This feature is, I believe, known in 
no other vertebrate. 

Text-fle-ure 10. 




FPr.Cor, 



Seijmouria hayloriensis Broili. — Outer surface of the incomplete riglit scai'ula 
aud precoracoid, X f . 

The fr.agment lying on the anterior end of the coracoid is clavicle. 

Eeference letters; — Cl., clavicle ; F.Gi., glenoid foranaen ; F.Pe.Coe., precoracoid 
foramen; F.S.Gi., supra-glenoid foramen ; Pe.Coe., precoracoid ; Sc, scapula. 



Shoiolder- girdle. — The shoukler-girdle has already been well 
described by Broili and Williston, but some features of its 
structure have so far escaped observation and others are worth)' 
of further emphasis. The scapula is exactly a,s Williston has 
described it, with a veiy broad supra-glenoid fossa, high up in 
which lies the small supra- glenoid foramen. My specimen shows 
clearly not only that it only ai'ticulates with a single coracoidal 
element, but also that it differs from the somewhat similar con- 
dition in . Varanoojjs in not presenting an articular face for a 
cartilaginous posterior coracoidal element. In fact the evidence 
makes it quite plain that there was only a single coracoid, 
corresponding to the anterior one of most Ootylosaurs, the 
posterior not being represented even by a cartilage. The pre- 
coracoid is as Williston has described it, with a very deep fossa, 
on its outer surface behind . the front of the glenoid cavity. A 
small glenoid and a much larger precoracoid foramen open into 
the fossa. 

The glenoid cavity is represented by a deep groove, V-shaped 
in section. This groove during life must have been filled up 
with cartilage, its margins map out a typical screw-shaped glenoid 
articulation. 

Sternum. — The hard matrix lying on the inner surface of the 



282 CAPr. D. M. S. WATSON ON SEYMOURIA, 

scapula and coracoid shows quite distinctly traces of an ossification 
or ossifications over an area about 2"5 cm. by 1'5 cm. This bone 
is of very open texture and has no definite surface, it was exposed 
so irregularly in development and its matrix is so extremely hard 
tha,t no account of its shape can be given. It is certain that it 
does not represent a. series of abdominal ribs or other dermal 
ossifications. It does not show any tendency to form long 
columns such as sternal ribs would be, and must apparently re- 
present a sternum, the first evidence of an. ossification in this 
element in a Lower Permian reptile. 

Text-fio'ure 11. 




Seymouria hayloriensis Broili. — Right side of the pelvis, X f. 
From vay skeleton. 

Pelvis. — The perfect pelvis of my skeleton agrees very well 
with Prof. Williston's figures and description, but the very 
well exposed ilia show certain significant differences from the 
young bone figured by Williston ('American Permian Yertebrates,' 
pi. xxix. fig. 7). In them tha,t part of the bone which lies 
behind the acetabulum and articulates with the ischium is quite 
large, and a more interesting diflerence is that the upper margin 
of the bone has a distinct projection towards its anterior end. 
This feature is shown by both right and left ilia. 

Femur. — The two well-preserved femora of my skeleton agree 
generally with Williston's figures, but appear to be considerably 
less massive. The lower end is very markedly divided into 
condyles. 

Tarsus. — Both hind limbs are in position with the femur 
directed forward with its head in the acetabulum. The knees 
are very strongly flexed. The right foot lies naturally articu- 
lated, but has been slightly laterally compressed, so as to slide 
the head of the third over that of the fourth metatarsal. Only 
four tarsals are preserved, and it appears extremely unlikely that 
any more were ossified at the death of the individual. The 
■ tarsus and foot are exposed from the plantar surface. 

There are three proximal tarsals all closely ai-ticulated with 
one another and with the tibia, but separated by an interval 
of about 8 mm. from the fibula. 



THE MOST PRIMITIVE KNOWN REPTILE. 283 

The fdbiale is a small squarish bone with rounded edges and 
corners. It has a somewhat definite edge towards the inter- 
medium. 

The intennecliam is a well-formed bone with a cylindrically 
concave ventral snrface. Towards the tibia and fibula it has 
well-foi-med edges, which are separated proximally by a smooth 
well-rounded notch of considerable breadth. It articulates with 
both tibiale and fibulare, tlie faces towards those bones being- 
separated distally by a small smooth notch. 

Text-fieure 12. 




Sei/mouria batjloriensis Broili. — Kiglit hind leg of my skeleton, X f . 

The bjiies are represented as thej^ lie in the matrix. The dotted part of the fibnla is 
exposed on the lower surface of the block and is represented as seen through 
it, and its distal end beyond the line which crosses it is drawn reversed from 
the bone of the left side. 

The fibulare is a small rounded bone resembling the tibiale in 
general characters. It is separated from the intermedinm by a 
definite layer of matrix. 

The only other tarsal preserved is a small rounded element 
above the upper end of the 3rd metatarsal. 

In text-fig. 12 1 have represented the right femur, tibia, fibnla, 
and proximal tarsals in the position in which they lie in the 
.matrix. The part of the fibula indicated in dotted lines is 
exposed on the other side of the block, and the distal strip of the 
fibula about 1 mm. wide is restored from the bone of the left 
side, a,s the right bone is slightly weathered here. 

In text-fig. 13 1 have restored the tarsus by moving up the 
fibula into contact with the proximal tarsals, the resulting 
accurncy of fit is good evidence for the reliability of the figure. 

This leg diflers from Prof. Williston's figure somewhat in the 
shape of the lower end of the fibula, and much more importantly 
in showing the clearest evidence of an intermedium. There is no 



284 



CAPT. D. M. S. WATSON ON SEYMOURIA, 



doubt that the distal notch in that bone formed part of the fora- 
men which occurs in all Permian reptiles between the "tibials" 
and fibulare. 

Text-fififure 13. 







SeyiHouria hayloriensis Broili. 

Right tibia, fibula, tarsus, and pes, reconstructed from my skeleton, 
the cross-hatched phalanges are not preserved. 

The left foot retains a complete 2nd toe and a third which has 
obviously been complete but from which part of the first and 
the whole of the second phalange are missing, the piece of 
matrix containing them not having been collected. These toes 
have respectively 3 and 4 phalanges, the terminal one being in 
each case a little conical bone. 



The foregoing description when read in connection with those 
given by previous authors renders Seytnouria nearly as well 
known as any reptile recent or fossil, and should allow of a very 
thorough study of its taxonomic relationships and of the bearing 
of its structure on morphological problems. Broili in his original 
discussion recognised the extraordinary Stegocephalian appear- 
ance of its skull. Williston, from a study of the whole skeleton, 
concluded " That the relationships of Seymouria are not very 
intimate with any other reptiles known from the Permian ; at 



THE MOST PRTMlTlVE KNOWN REPTILE. 



285 



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a a a p 
a ^ ^ cfl 

S 'S ^ 60 



+3 =S r^ TJ 

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p -M 03 ' — ' 

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286 CAPT. D. M. S. WATSOK^ ON SKYMOURIA, 

least any that are tolerably well known. In the skull the 
presence of the deep otic notch, the ai-i-angement and number of 
the temporal bones, the slender, elongate teeth, longest in the 
niaxillaj, and the shape, all differentiate the genus widely from 
either Diadectes or Limnoscelis, ns well as the chief forms now 
referred to the Pariotichidpe, and also from the known foreign 
forms." "In any event, it is certainly very remarkable tliat 
this cotylosaur reptile with all its other strange a.mphibian 
affinities should mimic so closely the temporal structure of the 
real amphibians." " However, it is very much of a question 
whether these resemblances [to Amphibia in skull and limbs] 
are so much the result of heredity and relationships as of 
adaptive, parallel, or convergent evolution. We have been 
speculating on the assumption that the known temnospondylous 
amphibians like Caco'ps, Eryops^ Euchirosaurus^ are, if not the 
actual ancestors of the reptiles, their first or second cousins. 
But this presumption is, in my opinion, quite unjustified." 

In several papers during the last five years, I have upheld the 
view that Seymouria is the most primitive of all known reptiles, 
and that its resemblances to Temnospondyls, particularly to the 
Embolomeri, are due to inheritance. I now propose in the light 
of a ps-actically complete knowledge of the skeleton in the 
Ootylosaurians Diadectes, Lahidosaurus, Captorhinus, Limnoscelis, 
Pariasmtrus, and Procolophon, and in the Temnospondylous 
Amphibia Eryops, Gacops, Tretnatops, T rimer or achis, Archego- 
saurus, Lydekkerina, and Bhinesicchus, and a good knowledge of 
the skull and much of the skeleton in many Embolomeri, to 
discuss the position of Seymouria in detail. 

Lahidosaurus and Captorh'mus belong to the same family ; 
each of the other forms' represents an independent family ; Dia- 
dectes, Paraisaicrics, and Procolojyhon represent the superfamily 
Diadectomorpha. The other Cotylosaurs are Captorhinomorphs 
(Watson, 1917). 

The skull and lower jaw of Seymouria as a whole resemble those 
of Labyi-inthodonts in their reticulate ornamentation. Similar 
sculpture is retained by the Captorhinomorpha alone amongst 
Cotylosaurs, that of other types being less markedly composed of 
pits, sometimes elongated to form long twisted channels. 

Basis Cranii. — The basioccipital of Seymouria agrees with those 
of all other Lower Permian Cotylosaurs, lachitomous Amphibia, 
and certain Embolomeri, in being excluded froin the brain-cavity 
by the meeting above it of special flanges from the exoccipitals. 
Certain Embolomeri (Pteroplax) have the basioccipital entering 
into the border of the foramen magnum, a condition which 
appears to be the primitive one for Tetrapods. Seymouria is 
unique amongst Cotylosaurs in the very large part which the 
exoccipitals play in the condjde. They form well-marked, down- 
wardly-projecting areas strongly reminiscent of those of Eryops, 
and not exactly paralleled in' any othei- reptile. 



l^HE Most primitive known reptile. ^87 

The large ventral exposure of the basioccipital is a reptilian 
character not achieved in Amphibia. 

Basisphenoid. — The presence of large powerful tubera basi- 
sphenoidales in the basis cranii of Seymouria is a reptilian 
character. Definite tubera do occur in certain Temnospondyls 
(Enjops), but these are always small. In Embolomeri tubera ai-e 
absent. 

The short basipterygoid processes of Seymouria supporting the 
pterygoid through the intervention of a special bone are unique, 
nothing similar occurring in any other known adult reptile or 
amphibian. The shape and position of the pterygoid render it 
certain that these special bones cannot be the epipterygoids, 
which in Dimetrodon are known to articulate with the basi- 
sphenoid. tSwinnerton and Howes, showed that in the develop- 
ment of the skull of Splienodon special articular cartilages are 
developed between the basipterygoid processes and the pterygoid, 
and it is not impossible that these are the representatives of the 
articular bones in Sey^nouria. 

• On the other hand, Gaupp has shown that in Lacerta the 
basipterygoid processes contain independent centres of ossification, 
and it is feasible and attractive to regard the Seymouria bones as 
permanently separate autogenous basipterygoid processes. 

The way in which the sides of the lower surface of the basi- 
sphenoid of Seymouria pass on directly into the outer surface of 
the prootic is reptilian, agreeing exactly with the conditions in 
most Cotylosaurs, The structure of this region is, however, 
equally similar to that of Pteroplax and another Middle Coal- 
Measure Embolomerous amphibian ; but it is quite difierent 
from the conditions in Rachitomi. 

The parasphenoid of Seymouria is of normal reptilian type, 
and also agrees with that of Embolomeri. It diff"ers in its small 
size from that of Rachitomi. 

Brain-case and Otic region. — The brain-case and otic region of 
Seymouria present a wonderful mixture of features occurring in 
other Cotylosaurs and in Labyrinthodontia. 

Seymouria is unique amongst reptiles in not possessing an 
ossified supraoccipital. It also diflfei-s in this respect from the 
Embolomeri, but agrees exactly with the conditions in most 
Rachitomi. 

The whole structure of the exoccipital, its relations to the 
basioccipital and paroccipital, and to the skidl-roof especially in 
its connection with a special descending lappet from the dermo- 
supraoccipital, is that of such an amphibian as Trim,erorachis, 
which resembles Seymouria even in the unusual position of the 
hypoglossal foramen. 

The Rachitomous amphibian in which the brain-case is most 
completely known is Eryops, where the structure has been 
described by v. Huene, Bi'oom, the present writer, and Williston. 

Although Prof. Williston criticises certain points of my 



288 CAPT. D. M. S. WATSON ON SKYMOURlA, 

description, the discrepancies between his account nnd mine seem 
to be ahiiost entirely, and in all important points are entirely, 
difi'erences of interpretation and not of structure. The only sig- 
nificant difference is that the foramen I hold to be for the Xtli 
nerve Williston takes to be that for the Xllth. It is undeniable 
that it does greatly resemble a hypoglossal foramen, but in the 
several specimens in the American Museum which show both its 
outer opening and the suture between the exoccipital and par- 
occipital, there is no doubt that it opens between these two bones 
as a vagus foramen always does. The course of the Xth nerve 
demanded by Williston's interpretation leads through the large 
cavity, which is definitely shown by an American Museum skull, 
which I have described and figured, to have housed the vestibule 
of the inner ear, and passes out high up in the lateral walls of 
the skull through a foramen which is certainly absent in certain 
well-preserved New York material, and therefore cannot be for 
any cranial nerve. I thus still prefer my own interpretation 
of the structure, although for theoretical reasons I should wish 
to see a Xllth nerve in Eryops as in so many other Labyrin- 
thodonts. 

The brain-case and otic region of Seymouria resemble the 
corresponding regions of a Rachitomous amphibian in the follovv- 
ing features : — 

1. The large opening from the brain-cavity to that for the 

inner ear. 

2. The position of the inner ear in the side-wall of the skull. 

3. The way in which the irnier anterior corner of the prootic 

reaches up to the skull-roof. 

4. The extremely massive paroccipital process. 

5. The upward clirection of the paroccipital process. 

6. The presence of an occipital flange from the tabular cover- 

ing a large area of the back of the paroccipital. 

The only difi"erence of importance between the ear-region of 
Seymouria and Rachitomous amphibia is that in the reptile the 
fenestra ovalis lies very low down, being bounded below by the 
basisphenoid and basioccipital along the posterior margin of the 
tubera basisphenoidalis, whilst in the Amphibia the lower border 
of the fenestra is formed by the parasphenoid, the basioccipital 
at any rate never entering into its margin. 

In characters 4, 5, 6, Seymouria differs from and is obviously 
more primitive than all other reptiles, for it is easy to derive 
the very diverse conditions in other Cotylosaurs fi'om those in it. 

The condition of the fenestra ovalis in Seymouria is paralleled 
amongst Cotylosauria only by the Captorhinidse, which in this 
region present an almost identical structure. 

The conditions in such types as Diadectes and Pariasaurus 
seem to depend on a reduction in size of the stapes and a corres- 
ponding diminution of the fenestra which receives its proximal 



THE MOST PRIMITIVE KNOWN REPTILE. 289 

end. The reduction seems to have taken place at the venti'al 
end of the opening. The Captorhinidfe retain a very large 
stapes. 

^phenet/wioid.^^— The anterior end of the brain of Seyviouria is 
surrounded by a badly known bone, which agrees exactly in its 
main features with the sphenethmoid of Eryops, and with the 
same bone in Fariasaihrus and similar bones in other Cotylosaurs. 

Palate. — The palate of Seymoti,7-ia is in essentials identical with 
that of the Embolomerous Labyvinthodonts. 

The resemblances depend on the identity in shape of the large 
pterygoid, meeting its fellow in the middle line anteriorly, 
laterally articulating with the prevomer, palatine, and ectoptery- 
goid, and having the quadrate ramus formed by a vertically 
standing plate reaching nearly up to the skull-roof, and bending 
round behind the quadrate to meet the squamosal in a long 
sutvire. 

This bone is covered, apparently all over, by a shagreen of 
granular teeth in Seymouria and Pteroplax. 

• The palatine of Seymouria agrees exactly with that of Temno- 
spondyli in structure, the most striking similarity lying in the 
tusk. This tooth is unique amongst reptiles in its mode of 
insertion in a pit Avhich also encloses the replacing tooth, but 
agrees exactly with all the large palatal teeth of Stegocephalia. 

The ectopterygoid of Seymoiiria agrees exactly with that of 
Eryops in contributing to a rudimentary flange which is applied 
to the lower jaw, and consists essentially of a deflected corner 
formed by pterygoid and transverse. The only feature of the 
palate of Seymouria which cannot'-be matched in Temnospondyli 
is the approximation of the posterior nares to the middle line. 

The palate of Seymouria differs from those of all other 
reptiles in the presence of the palatine tusks and the very weak 
development of the flanges which face the lower jaw. 

Tlie Captorhinid palate more nearly resembles that of Seymouria 
than those of other Ootylosauis, but differs in not having the 
quadrate ramus of the pterygoid reaching the squamosal, in the 
stronger transverse processes, and the loss of the palatine tusk. 
The maxillary teeth of Seymouria exactly resemble those of 
Temnospondyls in being fused to their base and to a labial wall 
of bone, in having fluted roots, and in being replaced alternately. 
No other Cotylosaur has at all a similar dentition. 

Prof. Broili has already shown that in its general build, in the 
deep otic notch, and the backwardly-inclined quadrate, and in the 
presence of the intei'temporal, the skull of Seymouria resembles 
that of the Temnosj)ondyls. In all these features it difiers from 
all other reptiles. 

The elongated lachrymal reaching from the orbit to the nostril 
is, however, a reptilian characteristic known in scarcely any 
Temnospondyl. 

The septomaxilla of Seymouria agrees with that of Temno- 



unique 


in 


the 


length 


of 


its 


clearly 


in 


the 


The ] 


pair 


■ of 



290 CAPT. D. M. S. WATSON ON SEYMOURIA, 

spondyli in lying within the nostril and not appearing in the 
face. The condition of this bone is known in few other Coty- 
losaurs. 

Loioer jaiu. — The mandible of Seymouria is typically Laby- 
rinthodont in structuie, agreeing very closely with that of 
Trimerorachis, in possessing a post-splenial and three coronoids. 

The presence of a patch of small granular teeth on the second 
coronoid is also a point of resemblance to most Kachitomi. 

The single large tusk on the 3rd coronoid is unique amongst 
Tetrapods, being unparalleled in either Amphibia or Reptilia, but 
of constant occurrence in Osteolepid fish. 

The non-fusion of the articular with the surangular is a point 
in which Seymouria differs from the Temnospondyls and agrees 
with the majority of reptiles. 

Vertebral column. — The atlas of Seymouria is 
lateral compression and great antero-posterior 
neural elements. This feature comes out most 
large spine composed of two apposed halves, 
neural elements together much more resemble a normal dorsal 
neural arch than in any other known Tetrapod, and are un- 
doubtedly extraordinarily primitive. The odontoid is thoroughly 
i^eptilian. * 

The axis of Seymouria in its complete unspecialisation, resem- 
bling as it does the vertebra next behind it, is quite different 
from that of any other reptile, whilst it agrees with that of 
Temnospondyls, where the 2nd and 3rd vertebi'te have a similar 
resemblance to one another. 

Prof. Williston in a recent paper has given an account of the 
mode of evolution of a Seymouria vertebra from an Embolo- 
merous type, with which I am in perfect agreement. He shows 
that the Seymouria vertebra is more primitive than that of any 
other known reptile in having a larger intercentrum, only 
slightly reduced from the disk which represents that bone in 
Embolomeri. 

The presence of distinct jirocesses for the head of the rib in 
certain Seymouria intercentra is a condition known in many 
Rachitomi and occurring in no other reptile. 

The presence of a single sacral vertebra only is an Amphibian 
feature very rare in reptiles. The wide separation of the centra 
of the caudal vertebrae suggesting the presence of a ring-shaped 
intercentrum, is a very primitive feature. 

On the other hand, the massive neural arches of the pre-sacrals 
with horizontally placed and widelj' separate zygapophyses are 
tj'pically Cotylosaurian, and occur in no Labyrinthodont what- 
soever. The caudal neural arches are also of typical reptilian 
type. 

Rihs.-r-^Ti possessing double-headed ribs throughout the whole 
pre-sacral part of the vertebral column, Seymouria differs from 
all other known Cotylosaurs and agrees with the Embolomeri. 
There can be no doubt that two-headed ribs are primitive, and 



THE MOST PRIMITIVE KNOWN REPTILE. 291 

that they are preserved on the atlas and axis in all early reptiles 
together with a permanent primitive temnospondylous structure. 

The double-headed caudal ribs of Seymouria are unique 
amongst Tetrapods. 

Shoulder- girdle. — I have endeavoured to show that primitively 
the Tetiapods had only a single cartilage element on each side of 
the shoulder-girdle, that a precoiucoid was then added to this, 
and subsequently a coracoid appeared. If this view be justified, 
Seymouria, which has only a precoracoid and presents no trace 
even of a cartilaginous coracoid, is primitive, more so indeed th.an 
any other known lower Permian reptile. 

The preservation of all three foramina of the Rachitomous 
shoulder-girdle is a primitive feature known in few other 
reptiles. 

Pelvis. — The ilium of Seymouria d^iWevs. in the antero-posterior 
elongation of its dorsal end from that of any Rachitomous 
Labyrinthodont, but in this chai-acter it exactly i-esembles the 
four known Embolomerous ilia. It agrees with these also in the 
pi'ocess from the upper edge toward the anterior end of that 
border. Several cotylosaur ilia resemble that of Seymouria in 
the production of the caudal end of the upper part of the bone. 

Fore Ihnh. —y^iWi^tow has already called attention to the 
remarkably Temnospondyl appearance of the humerus, but he 
has also pointed out that this bone differs from all amphibians, 
and resembles all early reptilian humeri in the presence of an 
entepicondylar foramen. 

Hind limb. — Williston has shown how much the Seymouria 
femur I'esembles that of such a Rachitomous amphibian as 
Eryops, and I have nothing to add to his account. 

It is, however, of great interest to note that it also presents a 
still more striking resemblance to the Lower Carboniferous 
(Lower Mississippian) femur which I recognised as probably 
I'eptilian and called Pajyposaurus traquari. This bone is very 
nearly as old as the oldest known amphibian bone, and far older 
than any other known reptile. 

The tarsus as revealed hy my skeleton is thoroughly amphibian 
in retaining a separate intermedium. In all other reptiles 
except Limnoscelis, where the fibulare is unossified, ancl the 
Plesiosaurs and Ichthyosaurs, there are onl}^ two proximal 
tirsals, the fibulare and a compound bone formed of the fused 
intermedium and tibiaJe. 

• The phalangeal formula is appaiently that common to all early 
reptiles. 



The foregoing series of comparisons show how wonderful a 
primitive and annectaiit form Seymouria is. In every part of its 
skeleton it shows a mixture of Temnospondyl and Reptilian 
characters, each recognisable, and in general showing little 
evidence of an intermediate condition. The whole eftect of its 



292 CAPT. D. M. S. WATSON ON SEYMOURIA, 

structure is that of a mosaic of separate details, some completely 
amphibian, some completely reptilian, and very few, if any, 
showing a passage leading from one to the other. In this fea- 
ture our study of Seymotiria lends support to the belief first 
upheld by the Mendelians and now accepted and used by many 
palaeontologists, that an individual animal consists of an aggre- 
gate of separate characters which are to a large extent capable of 
separate evolution, and may indeed proceed with their evolu- 
tionary change at very different relative rates in diflerent stocks, 
although of course they react on one another so as to produce a 
workable result. 

It thus appears desirable to have a tabular statement of the 
characters in which Seymouria shows an advance on Temno- 
spondyl structure, using so far as possible that of the Embolomeri 
as standard, and pointing out where these advances are in 
different directions from those which occur in the Rachitomous 
Labyrinthodonts, the cousins of the Reptilia, 

List of characters in which Seymouria has advanced 
above the Embolomeri : — 

1. The basioGcipital is depressed and no longer presents a 

round concave condyle. Rachitomi parallel. 

2. There are tubera basisphenoidales. Rachitomi parallel. 

3. The exoccipitals exclude the basioccipital from the brain- 

cavity. Some Embolomeri and Rachitomi parallel. 
[4. A hypoglossal foramen is present. ? If really absent in 
Embolomeri, Rachitomi parallel.] 

5. The exoccipitals reach up to be overlapped by the occi- 

pital flanges of the dermo-supraoccipitals. Rachitomi 
parallel. 

6. The supraoccipital is not ossified. Some Rachitomi 

parallel. 

7. The inner ear, though widely open to the brain-cavity, is 

distinctly separated by a rim of bone. Some Rachitomi 
' parallel. 

8. The fenestra ovalis is large and placed low down so that 

its lower margin is formed by that of the basipterygoid 
process. Rachitomi not parallel. 

9. The paroccipital is partly covered behind by an occipital 

flange from the tabular. Rachitomi parallel. 

10. The lachrymal extends from the orbit to the nostril. 

Rachitomi not parallel. 

1 1 . The ductus naso-lachrymalis lies in the substance of the 

lachrymal. Rachitomi parallel [Micropholis). 

12. The choanee lie near to the middle line. Rachitomi not 

parallel. 

13. The articulating surfaces of the zygapophyses are hori- 

zontally placed and the neural arches wide and swollen. 
Rachitomi not parallel. 



THE MOST PKIMITIVE KNOWN REPTILE. 293 

14. Intercentra with reduced and unossified dorsal halves 

occur. Rachitomi parallel. 

15. A distinct sacral vertebra is present. Rachitomi parallel. 

16. A cleithrum is absent. Rachitomi not parallel. 

17. An additional bone occurs on each side of the cartila- 

ginous shoulder-girdle. Some Rachitomi parallel. 

18. An entepicondylar foiamen is present in the humerus. 

Rachitomi not parallel. 

19. The presacral part of the vertebral column is shortened. 

20. An articular not fused with the surang-ular occurs. 

Rachitomi not parallel. 

21. The odontoid is a single bone. Rachitomi not parallel. 

The only characters which show that Seymouria is a reptile 
are : 8, 10, 12, 13, 16, 18, 20, 21, of the foregoing list. 

*It is specially to be noted that some of these [10, 18, 20] 
are features apparently of very minor interest, and that the 
really important characters, in the absence of knowledge of which 
no one would be justified in recognising the reptilian nature of 
'Seymouria, are those of the vertebral column. Nothing like the 
large swollen neural arches with horizontally placed articulations 
is known in Temnospondyls, and the small notochordal centra 
when taken in conjunction with the crescentic intercentra are 
equally distinctive. 

It thus appears that the vertebral column affords the best 
evidence for the reptilian nature of any Palaeozoic tetrapod. 

It is of interest to compare Seymouria in more detail with 
other Cotylosaurs and putative Cotylosaurs. 

Sauravits. — The small, rather imperfectly-known animal from 
the upper part of the Stephanian of France described by 
Thevenin as Sau7-avus costei was regarded by its describer as a 
reptile, and referred by Case to the Cotylosaviria. This attribu- 
tion has not yet been challenged ; it appears to have been 
founded on the presence in the hind foot of two proximal tarsals 
and a digital formula 2, 3, 4, ? ?. 

The structure of. the vertebral column shows beyond all doubt 
that this form is a Lepospondylous amphibian. There are no 
intercentra in any part of the column, and in the caudal region 
the haemal spines, which in all reptiles are supported by inter- 
centra (which in Mososaurs may fuse with the centra), project 
from the lower surface of the whole length of the " centrum." 
They have peculiar fluted ends, exactly similar to Ceraterj)eto7i 
and other types forming Miall's family IsTectridia. The dorsal 
''centra" ai^e slender hourglass-shaped bones shown by the 
section figured by Thevenin to be continuous with the neural 
arch. The presacral neural arches bear a flat lamellar expansion, 
the middle of which forms the rib-cai'rier, the whole strvicture 
being identical with that of a nectridian vertebra. In fact the 
whole column difiers so fundamentally from that of a Cotylosaur 

Proc. Zool. Soc— 1918, No. XXI. 21 



294 CAPT. D, M. S. WATSON OX SEYMOURIA, 

as to be incapable of ready derivation from an Embolomerous 
type, while it seems to resemble that of living Amphibia in 
being " pseudocentrous," i. e. with the apparent centrum derived 
from a membranous ossification lying between the notochordal 
sheath and the cartilaginous blocks of the embryonic vertebra. 
The animal is of considerable interest as showing that a reptilian 
type of foot may occur in Amphibia in no way connected with 
reptilian ancestry. 

Eosauravus. — The reniarkable skeleton from the Coal Measures 
of' Linton named by Williston Eosauravus copei, and described 
by Cope, Williston, Moodie, and Case, seems to be certainly a 
Cotylosaur if Case is correct in stating that the " neui'al arches 
are low and broad with horizontal zygapophysial faces and short, 
heavy spines." The " intercentral '*' position of the ribs, which 
has troubled certain American authors, really means very little. 
The skeleton (which I have not seen) is probably not wfell 
preserved, and if it be that of a reptile as primitive as SeymovTia, 
no doubt had double-headed ribs of which the capitulum articu- 
lated with the intercentrum. The slab may have been split so 
that only this head is preserved. 

The chief interest of the specimen lies in the proportions of 
the animal. Twenty-three left dorsal ribs are preserved, and the 
position of the left hand suggests that tw^enty-eight or more 
presacral vertebrte were present. There are preserved twenty- 
three caudals, the latter half of which series shows no signs of 
tapering, so that the tail may easily have been quite as long 
as the presacral part of the body. These proportions at once 
recall the aquatic Embolomeri. Pteroplax has more than 
28 presacrals, so has Pholidogaster, which animal has a tail as 
long as its body. The long slender form of Cricotus is familiar 
from Cope's two skeletons. Thus it is not improbable that 
Eosauravus retains a primitive form. 

In possessing only two proximal tarsals, it is undoubtedly 
more advanced than Seyniouria. 

Fapposaurus. — I have already pointed out that the Lower 
Carboniferous femur which is the type and only known material 
of Papposaurtcs traquari distinctly recalls ^eymouria. This 
resemblance increases the probability (admitted by Williston 
and Broili) that the form is a reptile. If so, it is by far the 
earliest known. 

Limnoscelis. — The large Cotylosaur from New Mexico described 
by Williston as Limnoscelis is in its limb-structure one of the 
most primitive known. The skull is, however, far more advanced 
than in Seymouria ; it retains no trace of the typical Eryopine 
shape of the latter skull, in particular the long slender otic 
notch of the more prirtiitive form is lost, the posterior surface 
appearing to be truncated. The occiput unfortunately has not 
yet been described in detail, but from Prof. Williston's ovit- 
line drawing it appears to be derivable from that of Seymouria 
by a migration downwards of the ends of the paroccipital 



THE MOST PRIMITTVE KNOWN REPTILE. 295 

processes, the fenestra ovalis retaining its position quite at the 
lower surface of the skull. 

The palate of Lininoscelis has advanced far beyond that of 
Seymouria in the wide interpterygoid vacuity and in the 
pronounced pterygo-transverse flanges. 

The lower jaw seems to have lost the post-splenial and pro- 
bably the anterior coronoids. 

The vertebrae in retaiaing a quite long spine are probably moi^e 
primitive than those of 8eymou7'ia, for all the Temnospondyls 
have high neural spines. 

The single-headed ribs are, however, an advanced feature. 

In the shoulder-girdle Limnoscelis is advanced in having a 
posterior coracoid element, but retains in its cleithrum a primitive 
bone lost by Seymouria. 

The pelvis of Limnoscelis much resembles that of Seymouria in 
the general form of the ventral surface, which is very reminiscent 
of Rachitomi in the short pubes. The posteriorly produced ilium 
is also a primitive feature retained by both reptiles. 

The hind leg of Limnoscelis resembles that of Seymouria in 
having a separate intermedium, but the conditions are diflFerent 
in the non-ossification of the tibiale in the former animal. 

Limnoscelis is thus in most ways' more advanced than Sey- 
mouria, although it is still a very primitive reptile. 

Diadectidoe. — The Diadectids, known from New Mexico, Texas, 
and probably also Europe {Stephenospondyhbs), are obviously 
much more advanced reptiles than Seymouria. They have a 
remarkably specialised palate and occiput — which region I have 
recently discussed in detail. The brain- case is theoretically 
derivable from that of Seymouria, but is so extremely modified 
that no useful end will be served by a further discussion. 

Pariasauridce. — With the Middle and Upper Permian group of 
the Pariasaurs, we pass to reptiles which have advanced much from 
the primitive Seymouria structure in all features of skull and 
skeleton. Some of these diflferences I have already pointed out, 
others are at once obvious from any comparison. One important 
feature which deserves emphasis is the high position of the 
fenestra ovalis, which lies on the front of the paroccipital process 
far removed from the basioccipital and the tuber basisphenoidalis. 
The opening is small and the stapes is a slender rod, without the 
much expanded base-plate it must have had in Seymouria. 

Procolophonidce. — Procolophon and its allies are the most 
advanced Cotylosaurs known. They differ extremely from Sey- , 
mouria in nearly all features. The differences which at joresent 
most interest me are those in the brain-case and otic region. In 
essentials these regions resemble the condition in Sphenodon, 
and differ from that in Seymouria in ways I have already pointed 
out, perhaps most markedly in the slender paroccipital process 
with a grooved lower surface in which the stapes lies, and in the 
high position of the fenestra ovalis. 

The Cotylosaurs so far considered all differ markedly from 

21* 



296 CAPT. D. M. S. WATSON ON SEYMOURIA, 

Seymouria in the condition of the brain-case and of the ear. 
Although their structure could in practically all points be derived 
from that of Seymouria, they do not in any way recall that 
reptile. In the Captorhinids we find features which distinctly 
hark back to the structure of Seymouria, although the group as 
a whole is one of the most advanced of the Texan Cotylosaurs. 

Through the work of Cope, Broili, Case, and especially "Willis- 
ton, we have a nearly perfect knowledge of the skeleton of both 
Captorhinus and the more advanced Labidosaiirus, and I have 
given a somewhat more detailed account of the occiput than is 
to be found in earlier writings. 

The skull of Captorhinus is advanced in the loss of the 
intertemporal and tabular and the very great reduction of the 
supratemporal ; it retains the primitive feature of a lachrymal 
reaching the nostril, and of a septomaxilla not appearing on 
the face. 

It resembles Seymoicria in the following important characters : — - 

1. The occipital condyle is tripartite, the exoccipitals forming 

a good deal of its surface. 

2. The fenestra ovalis is large, surrounded by the prootic, 

paroccipital, basioccipital, and basisphenoid ; it is 
placed low down in the skull so that its margin below 
lies on the border of the tuber basisphenoidalis. 

3. The basisphenoid is remarkably similar in the two 

genera. 

4. The palate of Captorhi'mis, though advanced in the loss 

of the palatine tusks and in the greater development 
of the ptery go-transverse flanges, is on the whole not 
unlike that of Seymouria. 

In the vertebral column the remarkably swollen neural arches 
of the presacral vertebrte and the short nearly obsolete neural 
spines, are points of resemblance betAveen the two forms. 

The Captorhinids have advanced over Seymoioria in the 
following ways : — 

1 . In the downgrowth of a process of the paroccipital between 

the lateral margins of the basioccipital and the inner 
border of the fenestra ovalis, so as to separate that 
opening more widely from its fellow. 

2. In the reduction of the paroccipital processes and the 

rotation of their ends downwai"ds. 

3. In an enormous widening of the supraoccipital. 

4. In a great deepening of the whole fcrain-case. 

5. In a reduction of the prootic so that it no longer reaches 

the skull-roof. 

6. In the loss of the tabular and the restriction of the 

dermo-supraoccipital to the occipital surface. 

7. In the loss of the intertemporal and in Lahidosaurus of 

the supratemporal. 



THE MOST PRIMITIVE KNOWN REPTILE. 297 

8. In the narrowing of the face. 

9. In the loss of the palatine tusks. 

10. In the better development of the ptery go-trans verse 

flanges. 

11. In the lateral compression of the quadrate so that its 

horizontal section is no longer U-shaped. 

12. In such reduction of the pterygoid that it no longer 

reaches the squamosal. 

13. In a considei^able compression of the lower part of the 

articular. 

14. In the loss of the two anterior coronoids and the post- 

splenial. 

15. In the somewhat more modified axis. 

16. In the presence of two sacral vertebrae. 

17. In the development of single-headed ribs. 

18. In the reduction of the intercentra. 

19. In the development of a coracoid in addition to the pre- 

coracoid. 

20. In the reduction and loss of ornament of the lower parts 

of the clavicles and the anterior end of the interclavicle. 

21. In the loss of the long posterior extension of the dorsal 

part of the ilium. 

22. In the more advanced humerus. 

23. In the loss of the long adductor crest in the femur. 

24. In the fusion of the intermedium and the tibiale. 

25. In the obliteration of the otic notch by the swinging 

backward of the upper end of the quadrate, so that the 
posterior end of the skull seems cleanly truncated. 

Thus the Captorhinids alone amongst Cotylosaurs present 
definite significant resemblances to Seyniouria, which in the case 
of the palate are due merely to the retention of a primitive 
structure, but in the important characters of the fenestra ovalis 
and the vertebrse are due to the possession by both types of a 
structure which is certainly not derived from amphibian ancestoi^s. 

The position and character of the fenestra ovalis common to 
Seymouria and Ca'ptorhinuH. is preserved throughout the whole 
series of mammal-like reptiles, the Pelycosaurs and all the later 
S. African types. This fact is clearly illustrated in the drawings 
of text-fig. 15, where the stapes is indicated by shading. 

This position of the fenestra is of course connected with that 
restriction of the inner ear to the lower part of the side wall of 
the brain-case wdiich I have shown to be one of the most constant 
and significant characters of the Anomodontia, and one which is 
entirely restricted to them and the Captorhinids, which Prof. 
Williston and the writer believe to be their ancestors. Com- 
parison of the occipital views of skulls in text-fig. 15, with similar 
drawings of any other reptiles, will bring out the diflerence 
between the two types of structure of this region clearly. 

Apart fi-om the Captorhinids, only Limnoscelis and Seymouria 



298 



CAl'T. L). M. S. WATSON UX SEYMOURIA, 

Text-figui-e 15. 









Series of occipital views, reduced to the same width, of the skulls of Aiiomodonts 
and of Cotylosaurs connected with that group. To show tlie low position, at 
the back of the tuber basisphenoidalis, which the fenestra ovalis occupies in 
these reptiles, its situation is indicated by the stapes. 



A. Seymouria bayloriensis.. 
C. Varanosaurus acutirostris. 
E. Diademodon broivni. 
Gr. Uimetrodon sp. 

A, from text-fig. 2 ; B, from Watson. 1916. 
in Munich; D, E, & H, from Watson, 1914; 
G, from a skull figured by Case, modified by other material. 

A-E form an approximate moi-phological series ; P & G represent an aberrant side 
line arising from C ; and H another distinct line also arising from a form near C. 



B. Captorhinus sp. 
D. Arctops willisto7ii. 
F. Deiopeus leptocephalus. 
H. Mormosaurus seeleyi. 

C. Restored from the type-skeleton 
F, modified from Watson, 1916; 



THE MOST PRIMITIVE KNOWN REPTILE. 299 

have this peculiar structure. I have ah^eady brigaded Livinoscelis 
with the Captorhinids. Seymouria is so much more primitive 
that it is remarkable to find in it a character of this kind 
connecting it clearly with a definite series of more advanced 
reptiles. 

It remains to be seen whether the other reptiles have lost this 
type of fenestra ovalis, or whether, as seems more probable, their 
ancestors never possessed it. 

It is a remarkably interesting fact that the primitive Pelyco- 
saurs such as Varanosaurus retain primitive features found in 
Seymouria which have been lost by the Captorhinids. Such 
are : — 

The retention of a supratemporal lost in Labidosaurus. 

The retention of a quadrate which is U-shaped in horizontal 

section. 
The retention of a suture between the pterygoid and squamosal 

behind the quadrate. 
The non-ossification of a posterior coracoidal element in 

Varoops. 
The retention of a considerably expanded lower end of the 

clavicle and a largely expanded interclavicle. 
The retention of a posterior process on the ilium. 

The characters in the above list represent features in which 
the Captorhinids are more advanced than the early Pelycosaurs ; 
they are, however, all acquired by later Anomodonts and represent 
parallel evolutionary changes which have gone on in the Capto- 
rhinids and Pelycosaurs independently since their separation. 
The most striking of these characters is the peculiar flattening 
and latei-al compression of the quadrate. In principle the 
quadrates of Caj)to7-hmus and Dimetrodon are identical, difiering 
only in the wide articular condyle of the Cotylosaur. As this 
type of quadrate is restricted to the members of the Superfamily 
Sphenacodontidse, we have a very remarkable case of parallel 
evolution, which in the original stock does not follow the main 
line of the evolution of the derived group, but duplicates a 
structure restricted to a single aberrant side line. 

The study of the Seymouria skeleton which fills this paper 
shows that that reptile is far more primitive than any other, 
presenting a strange mosaic of characters derived without change 
from the Embolomerous Labyrinthodonts Avith common reptilian 
characters marking distinct advance over an amphibian structure, 
and one or two which are restricted to the mammal-like reptiles 
and such Cotylosaurs as can be brought into connection with 
them. 

These facts have an important bearing on reptilian classifica- 
tion, suggesting as they do that all reptiles may be divided into 
two groups, one composed only of the Anomodontia and the 
Captorhinomorpha, the other of all other reptiles. 



300 CAPT. D. M. S. WATSOX ON SEYMOURIA, 

In the Cotylosaurs this division corresponds with that which 
I have previously founded on the character of the otic notch, 
and the mode in which a vertically placed quadrate is arrived at. 
I have recently found that this division on the character of the 
otic notch applies to all reptiles, affording a sharp distinction 
between the Anomodonts and all othei- groups, so that we have 
now two quite independent series of characters, the otic notch 
and the structure of the fenestra ovalis, which allow us to split all 
reptiles, Seymouria excepted, into two- groups. Seymouria itself 
shows chai-acters definitely connecting it with one of these. 

The curious way in which the structui-e of Seymouria is built 
up of perfectly well-developed amphibian characters and equally 
decisive reptilian features, those of intermediate type being very 
rare, afibrds a magnificent example of the way in which the 
evolution of great groups may liave taken place by the rapid 
change of all the definite morphological entities of which it 
may be regarded as made up, the chai:iges occurring quite inde- 
pendently and over a considerable time, the passage from the 
structure of the more primitive to the advanced group being quite 
gradual when viewed as a whole, but when further considered 
and analysed found to depend on a rapid evolution of separate 
regions apparently independently of each other. 

I hope that this study of '''feymouria will be regarded as placing 
beyond dispute the origin of the reptiles from the Embolomerous 
Labyrinthodonts. 

I wish to express my thanks to the Percy Sladen Trustees, who 
paid the expenses of my visit to Texas, to Prof. S. ^^^ Williston, 
who allowed me to study freely the superb collection of Lower 
Permian Tetrapods which he hasbniltup in Chicago, to President 
Osborn and the Staff of the American Museum for similar 
privileges there, and to Prof. F. Broili, who placed the treasures 
at Munich at my disposal. I regret that circumstances at 
present forbid my obtaining his permission to jDublish a descrip- 
tion of the Seymouria material of which he ga.ve so excellent a 
description. Finally, I wish to thank my wife for editing this 
paper and thereby removing many obscurities. 

Literature. 

Beoili, F. 1904. " Permische Stegocephalen und Reptilien aus 
Texas." Palseontographica, Bd. li. pp. 1-120. 

. 1908. " Ein montiertes Skelet A'on Lahidosaurus hamatus 

Cope, einen Cotylosaurier aus dem Perm von Texas." 
Zeitsch. deutsch. geol. Gesell. Bd. Ix. Heft 1, pp. 63-67. 

Broom, R. 1913. " Studies on the Permian Temnospondylous 
Stegocephalians of North Ameinca." Bull. Amer. Mus. 
Nat. Hist. vol. xxxii. pp. 563-595. 

Case, E. C. 1911. A Revision of the Cotylosauria of North 
America. 



THE MOST PRIMITIVE KNOWN REPTILE. 301 

Cope, E. D. 1896. " Second Contribution to the History of the 
Ootylosauria." Proc. Am. Phil. Soc. vol. xxxv, pp. 123- 

149; 

V. HuBNB, F. 1913, The Skull Elements of the Permian 

Tetrapoda in the American Museum of Natural History, 

New York. 
Thevenin, a. 1906. "Amphibians et reptiles du terrain houiller 

de France." Ann. de Paleontologie, tome i. pp. 145- 

163. 
WiLLiSTON, S. W. 1908. "The oldest known Reptile." Journ. 

Geol. vol. xvi. pp. 175-192. 
. 1910. " Desmospondylus anomalus." Bull. Geol. Soc. 

Amer. xxi. p. 280. 

. 1911. American Permian Vertebrates. Chicago, 1911. 

. 1916. "Synopsis of the American Permo-Carboniferous 

Tetrapoda." Contributions from the Walker Museum, 

vol. i. No., 9, pp. 193-236. 
^ , 1917. " Lahidoscmrus Cope, a Lower Permian Cotylosaur 

reptile from Texas." Journ. Geol. vol. xxv. pp. 309- 

321. 
. 1918. " (1) The Evolution of Yertebree." " (2) The 

Osteology of some American Permian Yertebrates, III." 

Contributions from the Walker Museum, vol. ii. pp. 75- 

112. 
Watson, D. M. S. 1912. " The Larger Coal -Measure Amphibia." 

Manch. Memoirs, vol. li. pp. 1-14. , 
. 1914. "Notes on Fa.r«?lo.sct^w■^^s aatitros^Hs Broili." Ann. 

k Mag. Nat. Hist. ser. 8, vol. xiii. pp. 297-310. 
. 1914. "The Deinocephalia, an Order of Mammal-like 

Reptiles." Proc. Zool. Soc. 1914, pp. 749-786. 
. 1914. "Notes on some Carnivorous Therapsids." Proc. 

Zool. Soc. 1914, pp. 1021-1038. 
, 1914. ^^ Procolo-phon trigonicejjs, a Cotylosaurian Reptile 

from South Africa." Pi-oc. Zool. Soc. 1914, pp. 735- 

747. 
. 1914. "On the Skull of a Pariasaurian reptile, and on 

the Relationship of that Type." Proc. Zool. Soc. 1914, 

pp. 155-180. 
, 1916. " On the Structure of the Brain-case in certain 

Lower Permian Tetrapods." Bull. Amer. Mus. Nat. 

Hist. vol. xxxv. pp. 611-636. 
. 1917. "A sketch Classification of the Pre-Jurassic Tetra- 

pod Yertebrates." Proc. Zool. Soc. 1917, pp. 167-186. 
. 1917. "The Evolution of the Tetrapod Shoulder-girdle 

and Fore Limb." Journ. of Anat. vol. Hi. pp. 1-63. 



ON THE FRESHWATER FISHES OF GREAT HRITAIN. 303 

EXHIBITIONS AND NOTICES. 

May 7th, 1918. 

Prof. E. W. MacBridb, D.Sc, F.R.S., Yice-President, 
in the Chair. 

The Ages of Elephants, as inferred from the Molar Teeth. 

Mr. R. I. PococK, F.R.S., F.Z.S., Curator of Mammals, ex- 
hibited a series of the molar teeth of Elephants that had died in 
the Gardens, and drew particular attention to the state of wear 
of the last tooth of an elephant known to be about 50 years old. 
Since this tooth comes into use in about the 40th year and had 
lost by wear more than one-third of its laminse in 10 years, 
Mr. Pocock concluded that the animal would have been toothless 
•and would have come to the end of her time before she was 70. 



Mr. E. Heron-Allen, F.L.S., F.R.M.S., F.Z.S., gave a lantern 
exhibition of Arenaceous Foraminifera of the e'enus Thurammina. 



May 28tli, 1918. 

Dr. S. F. Harmer, F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following Report on the Additions 
to the Society's Menagerie during the month of April 1918 : — 

The registered additions to the Society's Menagerie during the 
month of April were 209 in number. Of these 129 were acquired 
by presentation, 77 were received on deposit, and 3 were bred in 
the Menagerie. 

The following may be specially mentioned : — 

1 Barbary Sheep (Ammotragus lervia), born in the Menagerie 
ori April 18th. 

1 Blossom-headed Parrakeet (Palceornis cyanocephala) 6 , from 
India, presented by Miss H. F. Dunbar on April 27th. 



Mr. C. Tate Regan, M.A., F.R.S., F.Z.S., gave an account, 
illustrated by lantern-slides, of the Freshwater Fishes of Great 
Britain, with special reference to their value as food and to the 
possibilities of increasing their economic use. 



304 LT.-COL. S, M. COPEMAN 0\ A COLONY OF 

June 11th, 1918. 

A. Ezra, Esq., 7ice- President, iu the Chair. 

The Secretary read the following Report on the Additions 
to the Society's Menagerie during the month of May 1918 : — 

The registered additions to the Society's Menagerie during 
the month of May were 68 in number. Of these 14 were 
acquired by presentation, 46 were purchased, and 8 were bred in 
the Menagerie. 

The following may be specially pientioned : — 

2 Grecian Ibexes {Cajiva c(gagrus\, born in the Menagerie on 
May 9th and 15th. 

1 Ruby-throated Warbler [Calliope ccdliope), from India, pre- 
sented by W. H. St. Quintin, F.Z.S., on May 29th. 

1 Indian Chameleon (Chamceleon calcaratus), from Calabar, 
presented by A. M. Kinloch on May 11th; new to the Collection. 



Observations on a Colony of Burrowing Bees (Andrena fulva) 
on Primrose Hill. 

Lt.-Col. S. Moi\CKTON CoPEMAN, M.D,, F.R.S., F.Z.S., exhibited 
examples of the Burrowing Bee, Andrena fulva, and made the 
following remarks : — 

The existence of a flourishing colony of a burrowing bee 
{Andrena fulva) on the western slope of Primrose Hill, where, 
during the present year, it has spread over a much more extended 
area than previously occupied by it, appears worthy of record 
for the reason that one locality only in the neighbourhood of 
London — a small area on Hampstead Heath — is cited as a 
habitat of this insect in the British Museum Catologue of 
Hymenoptera. 

For the past six years this particular colony has been kept 
under observation, during which period the number of nests has 
not markedly vaiied from season to season until the present year. 
The unexampled spread of the colony this year (1918), curiously 
enough, was consequent on the manoeuvres of a Cadet Battalion 
last autumn having converted a previously grassy slope into a 
quagmire of mud, now represented by a smooth bare sui'face of 
clayey loam. This patch of ground the bees have found so much 
to their liking for nesting-purposes that (as indicated by a scale- 
plan drawn by my son last year) tliey have," in large measure, 
deserted their former haunts alongside a neighbouring path to 
seize upon the new territory so conveniently provided for them. 

Notwithstanding the hardness of the ground, the female bee 
excavates in it a tunnel which in many instances, on probing 
with a fine flexible wire, I have found to extend to a depth of 



BURROWING BEES ON PRIMROSE HILL. 305 

ten or twelve inches, and occasionally even more. This insect is 
usually about the size of the "worker" honey-bee, which she 
somewhat resembles in form. She is, however, handsomer, and 
also appears larger than her actual size owing to the abdomen 
especially being covered with a thick pubescence, which ranges in 
colour from a pale orange to a tawny-red in hue. This bee is, 
in consequence, a conspicuous object on a sunny morning as she 
ilies in mazy flight, usually a few inches only above the ground. 
The male insect is utterly unlike her, being smaller, less pubescent, 
and more sombre-toned in hue. 

As the outcome of six years' observations it has been found 
that over a period which has varied, according to prevalent 
meteorological conditions, from the middle of April to the first 
week in May the bees make their first appeai-ance quite suddenly, 
following on one or two days of warmer weather. For the first 
few days the number of males emerging greatly exceeds that of 
females, the relative divergence in numbers being as great as 
fifty to one. 

Careful examination of bare joatches of ground in the neigh- 
bourhood, so soon as the bees are first seen, discloses a number of 
small round holes which look as though they had been punched 
out with the point of a pencil. These are the open doorways of 
the burrows in which eggs were laid in the previous spring and 
in which the bees, passing through the stages of larva and pupa, 
have finally developed into the pei'fect insect. Herein the bee 
awaits the stimulus of warmth to give it strength to break 
through the slight covering of earth beneath which it has been 
sheltering since the previous autumn. 

Within a few days further, mating having taken place, the 
male bees are no longer met with, their brief life ending when 
the object of their existence has been attained. Almost imme- 
diately the females begin to dig^ their burrows, as indicated by the 
appearance of numerous little mounds like miniature volcanoes, 
eventually three or four inches in width and somewhat less in 
height. These are gradually built up of the tiny particles of 
earth thrown out, a few grains at a time, as the work progresses. 
In the first stages of the excavation the bee throws out the 
particles of earth between her hind legs, after the fashion of a 
dog when digging for a rat or rabbit, aiding the process by 
sweeping motions of her abdomen. Later on, however, when 
progress has been made beneath the surface, she brings up the 
grains of earth she has dug out, on the top of her head, generally 
waiting a few moments on reachi^ig the mouth of her burrow, 
before pitching them over the edge with a sudden jerk. The 
resulting mounds ai e easily rocognizable owing to the material 
of which they are composed being lighter in colour than the 
surrounding soil. 

On completion of the main burrow small nurseries are opened 
out from its sides, in each of which is deposited a carefully 
kneaded pellet of pollen and honey, which has to sufiice for the 



306 THE SECRETARY OX ADDITIONS TO THE MENAQERIK. 

needs of the giub which will develop from the egg laid upon 
its surface. This done, the little chamber is sealed up, and the 
process again repeated. 

During the numerous foraging expeditions necessary for obtain- 
ing sufficient pollen and nectnr for the sustenance of her future 
broods, no care is tnken to protect the burrow, a fact of which 
advantage is occasionally taken by a Noviada bee, too lazy herself 
to build nurseries or to gather stores of food. Waiting therefore 
till the rightful owner has set out on her travels, the " cuckoo " 
bee slips in and deposits an egg on any pellet of food of adequate 
size not yet sealed up. 

The Andrena bee not infrequently appears to experience con- 
sidei'able difficulty in recognizing her own burrow, as she may 
often be seen to tentatively explore the entrance to several before 
suddenly diving down her own. Marking the bee with a little 
powdered chalk as she leaves her burrow will render it easy to 
recognize the rightful owner on her return. 

The burrowing bee having finished her labours underground 
does not take the precaution of closing up the mouth of the 
burroAv, but on her death, within about a month from emergence 
from her winter quarters, leaves it to be graduallv obliterated by 
the levelling of the mound outside under the influence of rain 
and wind. 



October 22nd, 1918. 

Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following Report on the Additions 
made to the Society's Menagerie during the months of June, 
July, August, and September, 1918 : — 

June. 

The registered additions to the Society's Menagerie during the 
month of June were 38 in number. Of these 22 were acquired 
by presentation, 14 were bred in the Menagerie, and 2 were 
deposited. 

The following may be specially mentioned : — 

1 White-bearded Gnu {Connochcetes cdbojttbatus), born in the 
Menagerie, J"une 17th. 

4 Taurus Lizards {Lacerta taurica), new to the Collection, from 
Calamaria, Salonika, presented by G. H. Colt, F.R.C.S.,on Jvme 
20th. 

July. 

The registered additions to the Society's Menagerie during the 
month of July were 328 in number. Of these 24 were acquired 



ON WHEAT WEEVIL IN AUSTRALIA. 307 

by presentation, 9 were bred in the Menagerie, and 295 were 
deposited. 

The following may be specially mentioned : — 

1 Kiang [Equus kiaiig), from Tibet, born in the Menagerie, 
July 15th. 

1 Eland (Taicrotragus oryx)^ from South Africa, born in the 
Menagerie, July 16th. 

August. 

The registered additions to the Society's Menagerie during the 
month of August were 81 in number. Of these 59 were acquired 
by presentation, 9 were bred in the Menagerie, and 13 were 
deposited. 

The following may be specially mentioned : — 

1 Thar (^Hemitragus jemlaicus), from the Himalayas, bom in 
the Menagerie, August 10th. 

5 Coy pus [Myocastor coypus), from the Argentine, born in the 
Menagerie, August 16th. 

• 2 Verticillated Geckos (Gecco verticillatus), from Rangoon, pre- 
sented by Mrs. Hilda D. Sedgwick, August 10th. 

September. 

The registered additions to the Society's Menagerie during the 
month of September were 31 in number. Of these 15 were 
acquired by presentation, 2 were bred in the Menagerie, 8 were 
deposited, and 6 were received in exchange. 

The following may be specially mentioned : — 

Two hybrid Porcupines {Hystrix cristatus x H. africce australis), 
born in the Menagerie on September 20th. 



On behalf of Mr. E. Gerrard, Mr. R. I. PocoCK, F.R.S., 
exhibited the skin of an abnormally coloured red deer stag shot 
by Major R. C. Forster in Scotland, where it had been known 
for many years. It was a partial albino, the red hairs of the 
body being mixed with grey, the face, throat, and legs being 
white, the hoofs pale horn-colour, the muffle pink, and the eyes 
blue. 



Professor H. M. Lefroy, F.Z.S., Hon. Curator of Insects, gave 
an account, illustrated by lantern-slides, on the work he had 
accomplished for the Wheat Commission on Wheat Weevil in 
Australia. 



308 THU " NEW " IIABRTT DISEASE. 



November 5th, 1918. 

Prof. E. W. MacBride, D.Sc, F.R.S., Vice-President, 
in the Chair. 

Mr. D. Seth-Smith, F.Z.S., Curator of Birds, exhibited a 
mounted specimen of a hybrid Cockatoo ( d Roseate Cacatua 
roseicapiUa x $ Lesser Sulphur-Crested 0. stilphurea), bred in 
1917 at Hartwell House, Aylesbury, by Mrs. Lee. 



Diagnosis of Helminth Infections. 

Dr. R. T. Leiper, M.D., D.Sc, F.Z.S., gave a lantern 
exhibition on Diagnosis of Helminth Infections from the cha- 
racter of the eggs in the faeces. He stated that, by examination 
of the faeces of a living animal, the extent and specific nature of 
most helminthic infections could be accurately determined, and 
the method had been applied successfully as a routine practice 
in the case of man, rabbit, dog, cat, and pig, and was apparently 
capable of indefinite extension. 

The eggs of parasitic worms were constant in character and of 
great systematic importance. The ground-plan of the egg-shell 
indicated the genus or even subfamily to which the parasite 
belonged, and specific differences were found in slight but con- 
stant peculiarities in relative length and breadth, and in the 
conformation of excrescences on the surface of the shell. 



The '■'■ Neiv'''' Rabbit Disease. 

Dr. R. T. Leiper also gave a demonstration on the *' new '' 
rabbit disease. Examination of a large number of rabbits shows 
that the chief cause of mortality is a coccidial invasion of the 
intestinal wall or of the lining of the bile-ducts. According to 
Fantham and others the causal agent in both types of disease is 
Eimeria stiedce, bvit Dobell holds that the intestinal lesion is due 
to a distinct species. In many cases changes in the liver attri- 
buted to coccidiosis were the result of infection with Cysticercus 
pisiformis, the larval stage of the dog tapeworm Tcenia serrata. 
Large swellings in the region of the head and neck, suspected to 
be cancerous, were due to Ccenurihs serialis, the larva of the 
dog tapeworm Tcenia ccenitrus. Of relatively small economic 
importance are infections with the threadworm Oxyuris amhiguus 
and the tapeworm Ctenotcenia leuckarti. There is some evidence 
that a bacterial infection may occasionally be the cause of death. 

The coccidial infections pass from infected to healthy animals 
through the fseces. When freshly passed the coccidial oocysts 



THE SECRETARY OX ADDITIONS TO THE MENAGERIE. 309 

are not infective. They only become so after a period of delay, 
in which certain developmental changes take place. These 
changes proceed more rapidly in dry than in wet faeces. Pre- 
vention depends upon the systematic periodical removal and 
destruction by burning of all pellets and contaminated bedding, 
and the use of some fluid which will destroy such oocysts as 
remain in the hutch. 

Although several cases of coccidial infection in man have been 
recorded, Dobell maintains that in none of these cases is Eimeria 
stiedce tlie causal agent. There would appear therefore to be no 
risk of infection to man. 

The cystic stages of the tapeworms of the dog appear to occur 
chiefly in those rabbits fed with dandelions and other green 
stuffs collected from the roadsides, where the vegetation is 
especially liable to contamination with fgeces of dogs which have 
acquired their infections from eating uncooked rabbit offal. 



Professor H. M. Lefroy, F.Z.S., exhibited a series of lantern- 
slides, from photographs taken during the recent visit to the 
Zoological Gardens, Sydney, ISI. S.Wales. 



November 19th, 1918. 

Dr. A. Smith Woodward, LL.D., F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following Report on the Additions to 
the Society's Menagerie during the month of October, 1918: — ■ 

The registered additions to the Society's Menagerie during the 
month of October were 22 in number. Of these 8 were acquired 
by presentation, and 14 were deposited. 

The following may be specially mentioned : — 

1 Black-cheeked Ctircopitheque (CercGpithecus ascanms), from 
the Lower Congo, presented by Chevalier E. Carton de Wiart on 
October 2ud. 

2 Mississippi Alligators (Alligator mississip^jieusis), from North 
America, deposited on October 28th. 

1 Reticulated Python {Pytlion reticidatus), from the East 
Indies, deposited on October 11th. 

Special attention may also be directed to a captured German 
Carrier-Pigeon Loft, with 35 of the captured Pigeons, taken by 
the Canadians at Folies, Fi-ance, on August 9th, and presented 
to the Society on October 30th by the British Armies in France, 
at the suggestion of the War Office. 

Proc. Zool. Soc— 1918, No. XXII. 22 



310 CHIMPANZEE WITH PULMONARY TUBERCULOSIS. 

Miss K. Lanber, F.Z.S., described the method of preparing 
skeletons by the use of trypsin, and exhibited a number of 
successful examples from the Society's Prosectorium. 



Mr. E. Hatschek described his investigations into the forms 
assumed by drops and vortices of gelatii:i in various coagulants, 
exhibited a series of the forinations he had obtained which 
simulated animal structures, and demonstrated the method by 
which he obtained his results. 



Professor F. Wood-Jones, F.Z.S., exhibited a cast and a set of 
Rontgen-ray photographs taken from a Chimpanzee belonging 
to the Society, which had recently died from pulmonary tuber- 
culosis, and called attention to the possibility of diagnosing 
tubercle by this method in living subjects. 



Errata in Mr. L. A. Lanz's communication swpra^ pp. 11-17. 

Table, last line. 
For 26, read 36. 

Page 13. Footnote, line 3 from bottom. 
For 24, read 28. 

Page 14. Line 9 from bottom. 
For 37, read 39. 

Page 15. Last line. 

For 121, read 171. 



No. 184. 



ABSTRACT OF THE PROCEEDINGS 



OF THK 



ZOOLOGICAL SOCIETY OF LONDON, 

Octol)er 22nd, 1918. 



Dr. A. Smith Woodward, F.R.S., Yice- President, 
in the Chair. 



The Minutes of the last Scientific Meeting were confirmed, 

The Secretary read a list of additions to the Menagerie in the 
months of June, July, August, and September 1918. 

On behalf of Mr. E. Gerrard, Mr. R. I. PococK, F.R.S., 
exhibited the skin of an abnormally coloured red deer stag shot 
by Major R. C. Forster in Scotland, where it had been known 
for many years. It was a partial albino, the red hairs of the 
body being mixed with grey, the face, throat, and legs being 
white, the hoofs pale horn-colour, the muffle pink, and the eyes 
blue. 

Sir E. G. LoDER, Bart., Vice-President, read a communication 
entitled "Notes on the Beavers at Leonardslee, 1916-1918,"' 
containing evidence of the hitherto unrecorded fact that Beavers 
may breed twice in a season. 

A memoir by Mr. G. A. Boulenger, F.R.S., F.Z.S., on 

Madagascar Frogs of the Genus Mantidactylus, Blgr., was pre- 
sented to the Meeting. 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the 'Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillinys per annum, payable in advance. 



28 

Professor H. M. Lefroy, F.Z.S., Hon, Curator of Insects, gave 
an account, illustrated by lantern-slides, on the work he had 
accomplished for the Wheat Commission on Wheat Weevil in 
Austi-alia, 



The next Meeting of the Society for Scientific Business will be 
held on Tuesday, November 5th, 1918, at 5.30 p.m., when the 
following communications will be made : — 

Professor H. M. Lefrot, F.Z.S. 

Exhibition of Lantern-slides illustrating the Sydney Zoo- 
logical Gardens. 



Dr. R. T. Leiper, F.Z.S. 

(1) Lantern Exhibition on Diagnosis of Helminth Infections 
from the Character of the Eggs in the Ffeces. 

(2) Demonstration of the " New " Rabbit Disease. 

James F. Gemmill, M.A., D.Sc, M.D^RZ .S.^ 

On Ciliary Action in the Internal Cavities of the Cteno- 
phore, Pleitrobrachia pileus Fabr. 



The following Papers have been received : — 

D. M. S. Watson, F.Z.S., Capt. R.A.F. 

On Seymouria, the most primitive known Reptile. 

K. M. Smit h, A.R.C.S. 

A Comparative Study of certain Sense-organs in the Antenna 
and Palpi of Diptera. 



29 

Harold W . Leigh-Sharp, B.Sc. Lond. 

The Comparative Morphology of the Secondary Sexual 
Characters of Elasmobranch Fishes. The Olaspers, Clasper 
Siphons, and Clasper Glands. 



The Publication Committee desire to call the attention of 
those who propose to ofier Papers to the Societ}^ to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited as far as possible to the description of new results. 



• Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society of London, 

Eegent's Park, London, N.W. 8. 
October 22th, 1918. 



No. 185. 



ABSTRACT OF THE PROCEEDINGS 

OP THE 

ZOOLOGICAL SOCIETY OF LONDON. 

V November 5th, 1918. 

Professor E. W. MacBride, D.Sc, F.R.S., Vice-President, 
in the Chair. 



Mr. D. Seth-Smith, F.Z.S., Curator of Birds, exhibited a 
mounted specimen of a hybrid Cockatoo ( d Roseate dccatua 
roseicapiUa x $ Lesser Sulphur-Crested C. sulphurea), bred in 
1917 at Hartwell House, Aylesbury, by Mrs. Lee. 

The Chairman gave an account of a communication by 
Dr. James F. Gemmill, in which was described the cause of the 
ciliary action in the internal cavities of the Ctenophore, 
Fleurohrachia pileus. 

Dr. R. T. Leiper, M.D., D.Sc, F.Z.S., gave a lantern 
exhibition on Diagnosis of Helminth Infections from the chara- 
cter of the eggs in the faeces. He stated that, by examination 
of the faeces of a living animal, the extent and specific nature of 
most helminthic infections could be accurately determined, and 
the method had been applied successfully as a routine practice 
in the case of man, rabbity dog, cat, and pig, and was apparently 
capable of indefinite extension. 

The eggs of parasitic worms were constant in character and of 
great systematic importance. The groiind-plan of the egg-shell 
indicated the genus or even subfamily to which the parasite 



* This Abstract is published by the Society at its ofBces, Zoological Gardens, 
Rei'ent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issvied, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on 
the day of publication at the price of Sixpence, or, if desired, sent post-free 
for the sum of Six Shillings per annum, payable in advance. 



32 

belonged, and specific differences were found in slight but con- 
stant peculiarities in relative length and breadth, and in the 
confoi'mation of excrescences on the surface of the shell. 



Dr. R. T. Leiper also gave a demonstration on the " new " 
rabbit disease. Examina.tion of a large number of rabbits shows 
that the chief cause of mortality is a coccidial invasion of the 
intestinal wall or of the lining of the bile-ducts. According to 
Fantham and others the causal agent in both types of disease is 
Ewieria stiedce, but Dobell holds that the intestinal lesion is due 
to a distinct species. In many cases changes in the liver attri- 
buted to coccidiosis were the res\ilt of infection with Cysiicercus 
pisiformis, the larval stage of the dog tapeworm Tcenia serrata. 
Large swellings in the region of the head and neck, suspected to 
be cancerous, were due to Coenusus serialis, the larva of the 
dog tapeworm Tcerda ccenurus. Of relatively small economic 
importance are infections with the threadworm Oosyuris amhiguus 
and the tapeworm Ctenotcenia leucharti. There is some evidence 
that a bacterial infection may occasionally be the cause of death. 

The coccidial infections pass from infected to healthy animals 
through the fseees. When freshly passed the coccidial oocysts 
are not infective. They only become so after a period of delay 
in which certain developmental changes take place. These 
changes proceed more rapidly in dry than in wet faeces. Pre- 
vention depends upon the systematic periodical removal and 
destruction by burning of all pellets and contaminated bedding, 
and the use of some fluid which will destroy such oocysts as 
remain in the hutch. 

Although several cases of coccidial infection in man have been 
recorded, Dobell maintains that in none of these cases is Eimeria 
stiedce the causal agent. There would appear therefore to be no 
risk of infection to man. 

The cystic stages of the tapeworms of the dog appear to occur 
chiefly in those rabbits fed with dandelions and other green 
stuffs collected from the roadsides, whei-e the vegetation is 
especially liable to contamination with f feces of dogs which have 
acquired their infections from eating uncooked rabbit ofial. 



33 

The next Meeting of the Society for Scientific Business will be 
held on Tuesday, ISTovember 19th, 1918, at 5.30 p.m., when the 
following communications will be made : — 

The Secretary. 

Report on the Additions to the Society's Menagerie in the 
month of October 1918. 

Miss K. Lander, B.Sc, F.Z.S. 

Exhibition of Skeletons prepared by the "trypsin " method. 

E. Hatschek, 

Notes on Investigations into the Forms of Drops and 
Vortices of Gelatin in various Coagulants. 

D. M. S. W atson, D.Sc, F.Z.S. 

On Seymouria, the most primitive known Reptile. 



The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers siiould be condensed, and 
be limited so far as possible to the description of new results. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 

Zoological Society of London, 
Regent's Park, London, K.W. 8. 
November 12th, 1918. 



No. 186. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON.* 

November 19tli, 1918. 



Dr. A. Smith-Woodward, LL.D., F.R.S., Vice-President, 
in the Chair. 



The Secretary read a report on the Additions to the Society's 
Menagerie in the month of October 1918, calling special attention 
to a captured German Oarrier-pigeon Loft, with thirty-five 
of the captured Pigeons, taken by the Canadians at Folies, 
France, on August 9th, and presented to the Society by the 
War Office through Major A. H. Osman, Officer Commanding the 
English Carrier- Pigeon Service. 

Miss K. Lander, F.Z.S., described the method of preparing 
skeletons by the use of trypsin, and exhibited a number of 
successful examples from the Society's Prosectorium. 

Mr. E. Hatschek described his investigations into the forms 
assumed by drops and vortices of gelatin in various coagulants, 
exhibited a series of the formations he had obtained which 
simulated animal structures, and demonstrated the method by 
which he obtained his results. 

Professor F. Wood- Jones, F.Z.S., exhibited a east and a set of 
Rontgen-i-ay photographs taken from a Chimpanzee belonging 



* This Abstract is published by the Society at its offices, Zoological Gfirdens, 
Eegent's Park, N.W., ou the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Si.vpe?ice, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



36 

to the Society, Avliich had recently died from pulmonary tuber- 
culosis, and called attention to the possibility of diagnosing 
tubercle by this method in living subjects. 

Dr. D. M. S. "VVatson, F.Z.S., gave an account of a Memoir 
entitled " On Seymouria, the most primitive known Reptile," 
and illustrated his i-emarks by lantern-slides. 



The next Meeting for Scientific Business will be held on 
Tuesday, February 4th, 1919, at 5-30 p.m. Notice of the 
Communications to be made will be issued early in 1919. 



The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessaiy for the present that papers should be condensed and 
be limited so far as possible to the description of new results. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
ZooLoiGCAL Society of London, 

Regent's Paek, London, N.W. 8. 
November 2Qth, 1918. 



No. 187. 

ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON.* 

Felsruary 4th, 1919. 



Dr. S. F. Harmer, F.R.S., Vice-President, 
in the Chair. 



The Secretary read a report on the Additions to the Society^s 
Menagerie in the months of November and December, 1918. 

Mr. C. Da VIES Shebborn, F.Z.S., exhibited and made remarks 
on a letter written in 1693, by Malpighi to Dr. Mathew Faber. 

Sir Douglas Mawson read a communication on Australasian, 
Antarctic, and Subantarctic Life, and exhibited a large series of 
lantern-slides illustrating the scenery and mammals and birds 
of the South Polar Zone. He commented on the urgent need of 
international measures to preserve the fauna of these regions. 
The Chairman, expressing the sense of the Meeting, assured 
Sir Douglas Mawson of the active sympathy and support of the 
Zoological Society. 

A communication by Mr. R. I. Pocock was deferred to the 
next Meeting, with the consent of the Author. 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ' Proceedings,' free of extra charge, 
to allFellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



The next Meeting of the Society for Scientific Business will 
be held on Tuesday, February 18th, 1919, at 6.30 P.M., when the 
following communications will be made : — 

R. I. PococK, F.R.S., F.Z.S. 



On the External Chai'acters of the Existing Clievrotains. 
(Illustrated by lantern-slides.) 

K. M. Smith. 

A Comparative Study of certain Sense-Organs in the 
Antennae and Palpi of Diptera. (Illustrated by lantern- 
slides.) 

The following Papers have been received : — 

G. A. BouLENGER, F.R.S., F.Z.S. 

On a Collection of Fishes from Lake Tanganyika, with 
Descriptions of Three new Species. 

Miss Joan B. Pr octer, F.Z.S. 

On the Skull and Affinities of Rana suhsigillata, A. Dum. 



^ The Publication Committee desire to call the attention of 
those who propose to ofl'er Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

/Secretary. 
Zoological Society of London, 

Regent's Park, London, N.W. 8. 

February llth, 1919. 



No. 188. 



ABSTRACT OF THE PROCEEDINGS 

OP THE 

ZOOLOGICAL SOCIETY OF LONDON* 

February 18th, 1919. 

A. Smith Woodward, Esq., LL.D., F.E..S., Vice-President, 
ill the Cliair. 



The Secretary read a Report on the Additions to the Society's 
Menagerie in the month of January 1919. 



Mr. E.. I. PococK, F.R.S., F.Z.S., read a paper upon the 
external characters of existing Chevrotains (TraguHna), and 
showed that the Indian species, commonly cited as Tragidhis 
meminna, differs in so many important characters from the 
Malaysian species that it is necessary to sever it from them as 
a distinct genus, for which the name Moschiola, used by Thomas 
in a subgeneric sense, is available. In the absence of the inter- 
ramal scent-gland, in the structure of the penis, and in the 
retention of spots on the pelage, Moschiola is a more primitive 
type than Tragulus, and resembles the still more primitive West 
African genus Hyemoschus. 



Prof. H, Maxwell Lefroy, M.A., F.E.S., communicated a 
paper by Mr. K. M. Smith, on " A Compai-ative Study of certain 



* This Abstract is published by the Society at its offices, Zoological Gai-dens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with tlie ' Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 



Sense-Organs in the Antenna? and Palpi of Diptera" (illus- 
trated by lantern-slides). 



The next Meeting of the Society for Scientific Business wiU 
be held on Tuesday, March 4th, 1919, at 5.30 p.m., when the 
following communications will be made : — 

Dr. J. A. Murray, F.Z.S,, Acting Honorary Pathologist. 

Report on the Deaths in the Gardens during the Year 1918. 

G. A. BouLENQER, F.R.S., F.Z.S, 

On a Collection of Fishes from Lake Tanganyika, with 
Descriptions of Three new Species. 

Miss Joan B. Procter, F.Z.S. 

On the Skull and Affinities of liana subsigillata, A. Dum. 



The Publication Committee desire to call the attention of 
those who propose to offer Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited so far as possible to the description of new results. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
Zoological Society of London, 
Regent's Park, London, N.W. 8. 
February 2{)th, 121^. 



No. 189. 



ABSTRACT OF THE PROCEEDINGS 



OF THE 



ZOOLOGICAL SOCIETY OF LONDON/ 

March 4tli, 1919. 



Dr. A. Smith Woodward, F.R.S., Vice-President, 
in the Chair. 



Dr. J. A. Murray, F.Z.S., read a report on the deaths in the 
Gardens during the year 1918, iUustrating his remarks with 
lantern-slides. 

Mr. G. A. BouLENGER, F.R.S., F.Z.S., described a collection of 
Fishes from Lake Tanganyika, and drew attention to three new 
species. 

Miss Joan B. Procter, F.Z.S., gave a short account of her 
paper " On the Skull and Affinities of Rana subsigillata" drawing 
attention to several cranial characters either peculiar to this frog 
or held in common with R, adspersa, its nearest ally. 



* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the 'Proceedings,' free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on tha 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of iSix Shillings per annum, payable in advance. 



6 

The next Meeting of the Society for )Sciontific Business will 
be held on Tuesday, March 18th, 1919, at 5.30 p.m., when the 
following communications will be made : — 

H. R. A. Mallock, F.R.S., F.Z.S. 

"Some Points in Insect Mechanics." (Illustrated with 
lantern -slides.) 

F. Martin Duncan, F.R.M.S., F.R.P.S. 

Exhibition of Photographs and Lantern-slides of Marine 
Zoology. 

H. F. Blaauw, C.M.Z.S. 

" On the Breeding of Oryx gazella at Gooilust." 



The following Paper has been received : — 

Lancelot T. Hogben, B.A., B.Sc . 

" The Progressive Reduction of the Jugalin the Mammalia." 
(Communicated by Mr. H. W. Unthank,"B.A., B.Sc, F.Z.S.) 



The Publication Committee desire to call the attention of 
those who propose to ofier Papers to the Society, to the great 
increase in the cost of paper and printing. This will render it 
necessary for the present that papers should be condensed, and 
be limited as far as possible to the description of new results. 



Communications intended for the Scientific Meetings should 
be addressed to 

P. CHALMERS MITCHELL, 

Secretary. 
Zoological Society of London, 
Regent's Park, London, N.W. 8. 
March llth, 1919. 



Exhibitions and Notices (continued). 

Page 
The Secretary. Report on the Additions to the Society's Menagerie during the month 

of October, 1918 309 

Miss K. Lander, F.Z.S. Description and exhibition of the method of preparing skeletons 

by the use of trypsin 310 

Mr. E. Hatschek. Description and exhibition of investigations into the forms assumed by 

drops and vortices of gelatin in various coagulants 310 

Prof. F. Wood-Jones, F.Z.S. Exhibition of a cast and set of Rontgen-ray photographs 

taken from a Chimpanzee which had died from pulmonary tuberculosis 310 



PAPERS. 

10. 'Comparison between tlie Lower Jaws of the Cynodont Reptiles Gomphoffnathtts and 

Cynognathus. By Dr. Branislav Petronievics. (Text-figures 1-8.) 197 

11. On a new Genus of Extinct Mnseardine Rodent frona the Balearic Islands. By 

Dorothea M. A. Bate, Hon. M.B.O.U. (Plate I. and Text-figures 1 & 2.) 209 

12. A Case of Hermaphroditism in a Lizard, Lacerta viridis. By Noel Tayler, B.Sc. (Lond.). 

(From the Zoological Department, University of London, University College.) (Text- 
figures 1-3.) 223 

13. On two new Elasmobranch Fishes (CrossorMmcs Jurassicus, sp. nor., und Protospinax 

annectans, gen. et sp. nov.) from the Upper Jurassic Lithographic Stone of Bavaria. 

By Arthur Smith Woodward, LL.D., F.R.S., V.P.Z.S. (Plate I.) 231 

14. The Function of Pathological States in Evolution. By Morley Roberts 237 

15. Notes on the Eeavers at Leonardslee, 1916-1918. By Sir Edmund G. Loder, Bart., 

Vice-President Z.S 255 

lb. On the Madagascar Frogs of the Genus Mantidactylus Blgr. By G. A. Boulenger, 

F.R.S., F.Z.S 257 

17. Ciliary Action in the Internal Cavities of the Ctenophore Flmrohrachia pileus Fabr. 

By James F. Gemmil:,, M. A„ M.D., D Sc, F.Z.S. (Text-figures 1 & 2.) 263 

18. On Seymouria, the most primitive known reptile. By D. M. S. Watson, M.Sc, Capt. 

R.A.F., Lecturer in Vertebrate Paleontology, University College, London. (Text- 
figures 1-15.) - 267 

Titlepage i 

List of Council and Officers ii 

List of Contents iii 

Alphabetical List of Contributors vii 

Index • xi 



PLATES. 

1918, Parts III. & IV. (pp. 197-310), 



Plate Page 

Miss Bate : I. Hypnomys, gen. nov. et Leithia 209 



Woodward : I. 1 . Crossorhinus jiirassicus. 2, 3. Protospinax annec- \ , 
tans . . 



231 



NOTICE. 

The ' Proceedings ' for the jear are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. S. 1917, p. . . . The Distribution 
is usually as follows, but on account of war conditions Parts I. & II. were 
published simultaneously and Parts III. & IV. are similarly issued : — 

Part I. issued in March, 
„ II. ,, June. 

„ III. „ September. 

„ IV. „ December. 



'Proceedings,' 1918, Parts I. & II. (pp. 1-196), were published 
together on August 18th, 1918. 



The Abstracts of the ' Proceedings,' Nos. 184-189, are 
contained in this Part. 



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